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M. 


HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  1 


NOTES  ON  THE  AMERICAN  SOFT-SHELL  TURTLES 
WITH   SPECIAL   REFERENCE  TO  AM  YD  A  AGASSIZII 


By  Leoxhard  Stejxeger 


With  Thirty  Plates 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED  FOR  THE  MUSEUM 
May,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1 
have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  1 


NOTES  ON  THE  AMERICAN  SOFT-SHELL  TURTLES 
WITH   SPECIAL  REFERENCE  TO  AMYDA  AGASSIZII 


By  Leonhard  Stejneger 


With  Thirty  Plates 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED   FOR  THE  MUSEUM 

May,  1944 


PREFATORY  NOTE 

BY 

T.  Barbour 

As  has  been  well  known,  for  many,  many  years  before  his  death 
Doctor  Stejneger  was  engaged  in  the  preparation  of  a  general  treatise 
on  the  fresh  water  and  land  Testudinata  of  North  America.  After  his 
death  it  was  found  that  by  far  the  greatest  part  of  his  notes  were  in  the 
fragmentary  state  that  often  holds  in  an  early  stage  of  preparation 
for  publication.  The  only  section  which  might  be  said  to  be  in  pub- 
lishable  form  and  to  reflect  the  mature  conclusions  of  the  author  was 
this  part  comprising  his  summary  of  the  genus  Amyda.  For  many  and 
obvious  reasons  it  seems  desirable  to  bring  this  to  public  attention  and 
by  the  same  token  it  seems  presumptuous  to  change  it. 

The  Museum  of  Comparative  Zoology  has  the  good  fortune  to 
present  to  the  public  what  will  perhaps  be  the  last  example  of  the  work 
of  this  distinguished  veteran  worker  in  our  field.  By  the  consent  of  the 
authorities  of  the  United  States  National  Museum  this  institution  has 
permission  to  publish  this  work,  a  tribute  of  honor  and  respect  to  the 
author,  being  published  here  only  because  the  situation  induced  by 
the  state  of  war  makes  it  inconvenient  for  the  Government  Printing 
Office  to  handle  the  work  at  this  time. 


No.  1. — Notes  on  the  American  Soft-Shell  Turtles  ivith  Special  Reference 

to  Amy  da  agassizii1 

By  Leonhard  Stejneger 

INTRODUCTION 

The  number  of  living  indigenous  soft-shell  turtles  of  the  family 
Trionychidae  in  North  America  is  less  than  a  half  dozen,  while  more 
than  a  half  hundred  fossil  forms,  the  earliest  dating  as  far  back  as  the 
Cretaceous,  have  been  described  by  paleontologists  from  the  same 
region  which  embraces  the  habitable  part  of  North  America  east  of 
the  Rocky  Mountains  and  northern  Mexico. 

The  living  American  forms  have  been  considered  from  time  to  time 
by  prominent  herpetologists  as  belonging  to  several  genera  under 
varying  names,  according  to  their  views  on  the  genotypes.  Thus 
Agassiz  recognized  three  genera  and  so  did  Baur  and  Hay;  Cope  and 
True  reduced  the  generic  subdivisions  to  two.  Boulenger,  Siebenrock, 
and  recent  systematists  have  regarded  them  as  congeneric;  the  name 
has  varied  accordingly. 

NOMENCLATURE 

In  the  early  stages  of  systematic  herpetology,  before  the  idea  of  the 
"genotype"  had  become  generally  accepted,  there  was  no  fixed  rule 
which  would  designate  the  type  of  genera  with  more  than  one  species. 
A  sort  of  "selection  by  the  first  reviser"  was  early  practiced.  Thus  if 
an  author  subsequent  to  the  creator  of  a  plurispecific  genus,  in  sub- 
dividing the  latter,  retained  this  name  for  a  single  species,  applying 
one  or  more  new  names  to  the  rest  of  the  species,  his  selection,  as  a 
rule,  was  respected,  and  thus  the  idea  of  type  selection  by  "elimina- 
tion" became  a  more  or  less  general  practice.  But  because  of  the  un- 
willingness or  ignorance  of  some  workers  and  the  varying  and  uncer- 
tain application  of  this  method,  great  confusion  in  the  nomenclature  of 
genera  resulted.  This  is  what  happened  to  Geoffroy's  generic  term 
Trionyx  (1809),  which  has  been  the  cause  of  so  much  confusion.  In 
1816  Oken  was  the  first  one  to  subdivide  the  plurispecific  genus,  re- 
serving as  he  did  the  name  Trionyx  for  the  single  species  T.  granosa, 
leaving  the  others  under  the  name  Amy  da.  In  1830  Wagler  reported 
Oken's  monotypic  selection  of  Trionyx  granosa,  but  substituted  As- 
pidonectes  for  Amyda.   The  confusion  was  caused  by  Gray  who  in  his 

1  Published  with  the  aid  of  a  special  gift  from  Mr.  G.  R.  Agassiz. 


6  bulletin:  museum  of  comparative  zoology 

Synopsis  Reptilium;  1831,  erected  the  monotypic  genus  Emyda  for 
the  same  species.  He  did  so  in  ignorance  of  Wagler's  action  and  after 
seeing  Wagler's  work  called  attention  to  it  in  the  "Additions  and  Cor- 
rections" in  the  same  work,  p.  78.  Yet  in  the  "Catalogue"  of  1844  and 
his  later  writings  he  still  retained  Emyda.  Gray  was  followed  by 
Boulenger  and  many  writers  since  1889,  in  spite  of  Baur  and  Hay  who 
in  1898  and  1904,  like  Bonaparte  in  18361  and  1857'2,  upheld  Wagler's 
action. 

Following  these  dissertations  by  Baur  and  Hay,  the  case  of  the 
genotypes  of  the  genera  Amy  da  and  Trionyx  was  discussed  by  me  in 
1905  (Science,  new  ser.,  vol.  25,  Feb.  10,  1905,  pp.  228-229),  conse- 
quently before  the  adoption  (1908)  of  the  present  wording  of  Article 
30  of  the  International  Rules  of  Zoological  Nomenclature.  In  1905  the 
argument  was  based  on  the  process  of  elimination,  which  at  that  time 
was  considered  the  legitimate  process.  On  that  occasion,  the  type  of 
Trionyx  was  decided  to  have  been  selected  in  1816  by  Oken  as  first 
revisor  by  segregating  Trionyx  granosus,  supposed  to  be  synonymous 
with  Geoffroy's  T.  coromandelicus,  as  the  sole  species  in  the  restricted 
genus. 

However,  the  adoption  in  1908  of  the  changed  form  of  Article  30 
apparently  nullified  previously  accepted  type  selections  unless  they 
were  couched  in  the  stringent  language  of  the  new  version. 

The  new  Article  30  was  framed  and  phrased  for  the  purpose  of  doing 
away  with  the  uncertainties  of  personal  interpretation  unavoidable  in 
the  process  of  elimination.  It  was  intended  that  the  new  rule  should  be 
applied  literally;  hence  the  first  article  (I  a)  reads:  "When  in  the  origi- 
nal publication  of  a  genus,  one  of  the  species  is  definitely  designated  as 
type,  this  species  shall  be  accepted  as  type,  regardless  of  any  other  con- 
siderations." (Italics  mine.)  To  emphasize  this  intention  the  last 
paragraph  of  the  article  was  inserted  as  follows:  "The  meaning  of  the 
expression  'select  the  type'  is  to  be  rigidly  construed.  Mention  of  a 
species  as  an  illustration  or  example  of  a  genus  does  not  constitute  a 
selection  of  a  type."    (Italics  mine.) 

Applied  to  the  case  in  hand  the  facts  are  these: 

In  the  original  publication  of  Trionyx  by  Geoffroy  in  1809  (Ann. 
Mus.  Hist.  Nat.,  Paris,  14,  pp.  1-20,  pis.  1-5)  none  of  the  many  species 
was  definitely  designated  as  type.   Not  in  the  body  of  the  text  proper, 

1  Cheloniorum  Tabula  Analytica,  p.  8:  "22  AMYDA,  Schweigger  (Aspidonectes  WAGL. 
Trionyx  GRAY.  BELL.  Gymnopus,  DUM."  "23.  TRIONYX  WAGL.  {Emyda,  GRAY. 
BELL.  Cryptopas  DUM.)" 

2  Contrib,  Nat.  Hist.  United  States,  1,  p. 330:  "Trionyx  proper in  contradistinction 

to  Aspidonectes";  see  also  pp.  394-395. 


stejneger:  American  soft-shell  turtles  7 

but  in  the  Explication  des  planches  {explanation,  of  the  plates)  at  the  end 
of  the  article,  occurs,  however,  the  following  mention  of  the  trionyx 
of  Egypt  as  an  illustration  and  example  of  the  genus,  which  was 
omitted  in  the  simultaneous  reprint  of  the  article  in  the  Bulletin  of  the 
Philomatic  Society  of  Paris : 

"Le  trionyx  d'Egypte,  represente  planche  I,  vue  en  dessous  el  de  cote, 
nous  donnant  une  idee  exacte  du  port  et  des  caracteres  generiques  des 
trionyx,  nous  sommes  bornes,  dans  les  planches  suivantes,  a  faire 
figurer  les  seules  parties  caracteristiques  des  autres  especes,  telles  que 
leurs  carapaces  en  a,  et  leurs  plastrons  en  B."  l 

Even  if  this  kind  of  mention  as  an  "illustration  and  example"  may 
be  characterized  as  a  "clear  intention"  it  is  certainly  not  a  definite  desig- 
nation rigidly  construed. 

Consequently,  Geoffroy  having  failed  to  definitely  designate  T. 
aegyptiacus  as  the  type  of  Trionyx,  it  was  left  to  the  first  subsequent 
author  to  select  the  type,  and  that  was  done  by  Fitzinger  in  1843 
(Syst.  Rept.,  p.  30)  who  selected  T.  coromandelicus  Geoffroy,2  "and 
such  designation  is  not  subject  to  change." 

Having  now  disposed  of  the  type  designation  of  Trionyx  for  a  genus3 
in  the  subfamily  Cyclanorbinae,  the  question  of  the  status  and  type  of 
Amyda  arises. 

Among  the  new  species  described  by  Geoffroy  in  the  same  paper  of 
1809,  is  the  Trionyx  javanicus  (Ann.,  p.  15).  After  a  brief  diagnosis  he 
added  the  following  svnonvm: 

"Amyda  javanica  SCHNYEIGGER,  dans  un  manuscrit  communique 
a  l'Institut." 

This  publication  of  Schweigger's  monotypic  generic  name  Amyda 
clearly  establishes  its  availability  for  the  species  congeneric  with  his 
Amyda  javanica,  the  description  of  which  by  Geoffroy  is  regarded  as 
identical  with  Boddaert's  Testudo  cartilaginea  of  1770. 

Fitzinger's  use  (1835)  of  the  name  Amyda  for  a  section  (  =  subgenus) 
with  Trionyx  subplamis  as  type,  and  Agassiz's  subsequent  (1857)  ap- 
plication of  the  same  name  for  another  genus  with  Amyda  mntica  as 
type  are  consequently  both  void.  Dogania  is  the  proper  name  for  the 
former;  Euamyda  is  now  available  for  A.  mntica  as  a  generic  or  sub- 
generic  name. 

1  The  Egyptian  trionyx,  represented  on  pi.  1,  viewed  from  below  and  from  the  side,  gives  us  an 
exact  idea  of  the  aspect  and  the  generic  characters  of  the  trionyxes;  we  limit  ourselves  in  the 
following  plates  to  figure  only  the  parts  characteristic  of  the  other  species,  such  as  their  carapaces 
in  a  and  their  plastrons  in  b. 

2  Trionyx  coromandelicus  Geoffroy,  1809,  is  a  synonym  of  Testudo  granosa  Schoepff,  1792 

3  Synonyms:  Emyda  Gray,  1831  (not  of  Rafinesque,  1815);  Lissemys  Malcolm  Smith,  1931. 


8  bulletin:  museum  of  comparative  zoology 


Genus  Amyda1  Schweigger 

1809.  Amyda  SCHWEIGGER  in  Geoffroy,  Ann.  Mus.  Hist.  Nat.,  Paris,  14, 
p.  1  (monotype  Amyda  javanicus  =  T.  cartilagineus). 

1816.  Amyda  OKEN,  Lehrb.  Zool.,  2,  p.  348  (type  designated  by  Stejneger, 
1907,  Trionyx  euphraticus). 

1830.  Aspidonectes  WAGLER,  Nat.  Syst.  Amphib.,  p.  134  (type  designated 

1843  by  Fitzinger,  Trionyx  aegyptiacus  =  T.  triunguis). 

1831.  Trionyx  GRAY,  Synops.  Rept.,  p.  45  (type  T.  ferox)  (not  of  Oken, 

1816;  Wagner  1830). 

1835.  Gymnopus  DUMERIL  and  BIBRON,  Erpet.  Gen.,  2,  p.  472  (substi- 
tute name  for  Aspidonectes  Wagler)  (p.  484  Gymnopodus,  laps, 
calam.) 

1835.  Platypeltis  FITZINGER,  Ann.  Wien  Mus.,  1,  pp.  120,  127  (type  desig- 
nated by  Fitzinger  1843,  Platypeltis  ferox.) 

1835.  Pelodiscus  FITZINGER,  Ann.  Wien  Mus.,  1,  pp.  120,  127  (type  desig- 
nated by  Fitzinger  1843,  P.  sine?isis.) 

1842.  Aspedonectes    HOLBROOK,    North    Amer.    Herp.    2    Ed,   2,   p.    18 

(emendation). 

1843.  Potamochelys  FITZINGER,  Syst.  Rept.,  p.  30  (monotype  Aspidonectes 

javanicus). 

1844.  Tyrse  GRAY,  Cat.  Tort.  Brit.  Mus.,  p.  47  (type  T.  nilotica  =  T.  triun- 

guis). 

1856.  Tryonix  SAGER,  Peninsular  Journ.  Medic  Coll.  Sci.,  3,  no.  8,  Feb.  1856, 

p.  361  (emendation). 

1857.  Amyda  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  1,  p.  398  (mono- 

type A.  mutica)  (not  of  Schweigger). 

1864.  Rafetus  GRAY,  Proc.  Zool.  Soc.  London,  May  1864,  p.  81  (monotype 
T.  euphraticus). 

1864.  Aspilus  GRAY,  Proc.  Zool.  Soc.  London,  1864  (p.  83)  (type  Aspilus 
cariniferus). 

1869.  Callinia  GRAY,  Proc.  Zool.  Soc.  London,  1869,  p.  221  (type,  T. 
spiniferus). 

1895.  Platyrettis  KIRSCH,  Bull.  U.  S.  Fish  Comm.  1894,  p.  333;  typogr. 
err.?  for  Platypeltis). 

1900.  Aspidonectus  BEYER,  Proc.  Louisiana  Soc.  Nat.,  1897-1899,  p.  43 
(emendation). 

1  Name  of  uncertain  origin,  but  apparently  a  variant  of  Emys,  a  river  turtle.  In  the  synonymy, 
the  numerous  generic  terms  based  on  exclusively  Old  World  species  have  been  omitted.  Refer- 
ence to  these  may  be  found  in  Bulletin  United  States  National  Museum  No.  58,  1907,  p.  514. 


stejneger:  American  soft-shell  turtles  9 

The  generally  accepted  five  species  of  the  wider  genus  Amy  da  in 
North  America  naturally  fall  into  three  groups. 

1.  The  A.  mutica  group,  by  Agassiz  considered  a  distinct  genus  and 
by  many  accepted  as  a  subgenus  (for  which  the  name  Eliamyda 
may  be  substituted  as  Amy  da  Agassiz  is  preoccupied). 

2.  The  A.  feroz  group  including  the  species  A.  ferox,  spinifera  and 
emoryi. 

3.  The  A.  agassizii  group  containing  only  one  species  on  this  conti- 
nent. 

The  external  appearance  of  these  turtles  is  so  much  alike  that  the 
early  naturalists  had  difficulty  in  diagnosing  them  properly,  with  the 
result  that  their  taxonomic  history  is  full  of  misidentifications  and  mis- 
conceptions, even  to  the  extent  that  the  first  group,  at  least  on  one 
occasion,  has  been  suspected  of  being  only  the  sexual  form  of  one  of  the 
species  of  the  second  group. 

As  in  so  many  of  the  Old  World  soft-shelled  turtles  the  essential 
characters  are  recognizable  only  in  the  bony  structure,  so  also  in  our 
American  species.  The  most  important  ones  are  found  in  the  skull  and 
the  plastral  bones  and  they  will  form  the  main  subject  of  the  following 
discussion. 

Skull  Characters 

The  relative  proportions  of  the  skulls,  their  component  bones  and 
the  size  and  shape  of  the  various  fossae  and  foramina  vary  enormously 
with  age,  hence  comparisons  have  to  be  made  between  specimens  of 
about  the  same  size.  The  individual  variability,  which  is  considerable, 
increases  with  age  very  often  to  such  a  degree, — for  instance  the 
enormous  expansion  of  the  alveolar  surfaces  in  the  old  specimens  of 
A.  ferox, — as  to  make  even  group  definition  difficult.  Measurements 
of  a  large  number  of  skulls  have  therefore  to  be  made,  and  as  skulls  of 
approximately  equal  size  are  rarely  to  be  had,  the  measurements  have 
to  be  reduced  to  percentages  of  some  standard  dimension.  As  such  I 
have  selected  the  basicranial  length  (posterior  edge  of  occipital  con- 
dyle to  tip  of  snout).  The  analysis  of  numerous  measurements  has 
convinced  me  that  skulls  with  a  basicranial  length  between  40  and  70 
mm.  (average  about  55  mm.)  practically  represent  the  normal  propor- 
tions of  important  cranial  dimensions  of  our  American  trionychids. 
The  relative  proportions  of  the  various  parts  have  at  that  size  reached 
a  sufficient  stability  and  the  figures  consequently  are  comparable  inter 
se.  Unfortunately,  series  of  skulls  of  soft-shelled  turtles  are  not  numer- 


10  bulletin:  museum  of  comparative  zoology 

ous  in  museums,  especially  when  reduced  to  specimens  with  a  basi- 
cranial  length  between  40  and  70  millimeters,  hence  the  tables  pre- 
sented below,  based  as  they  are  exclusively  on  United  States  National 
Museum  material,  total  only  41  specimens;  nevertheless,  the  figures 
are  believed  to  be  fairly  representative. 

An  inspection  of  table  1  will  show  that  Amy  da  agassizii  possesses 
smaller  internal  choanae  (ch,  plate  1,  fig.  2)  than  the  other  species,  but 
a  slightly  larger  intermaxillary  foramen  (int.  max.  for.,  plate  1,  fig. 
2).  Were  it  not  for  the  latter  fact  the  distance  between  the  inter- 
maxillary foramen  and  the  choanae  would  have  been  greater  than  it 
is,  (viz.  12.5  against  only  7.7  to  9.7)  in  the  other  species;  in  other 
words,  in  A.  agassizii  the  longitudinal  diameter  of  the  choanae  equals 
their  distance  from  the  intermaxillary  foramen,  while  in  the  others  it 
is  much  greater,  in  A.  ferox  even  averaging  twice  as  much. 

The  table  further  shows  the  greater  width  of  the  alveolar  surface  of 
A.  agassizii  at  the  intermediary  age  of  these  specimens,  but  older  males 
(over  70  mm.  basicranial  length)  of  A.  ferox  acquire  an  increasing 
width  of  the  alveolar  surfaces  far  exceeding  in  proportion  even  that  of 
A.  agassizii  (plate  30).  The  width  of  the  alveolar  surface  of  the 
mandible  in  A.  agassizii  is  twice,  or  nearly  twice,  as  wide  as  in  the 
others  of  the  corresponding  size. 

On  the  underside  of  the  skull  A.  agassizii,  in  addition  to  the  small 
size  of  the  choanae  and  the  greater  width  of  the  alveolar  surface,  is 
characterized  by  the  position  of  the  suture  between  the  palatines  and 
the  basisphenoid  relative  to  the  posterior  edge  of  the  temporal  fossa. 
This  is  coincident  with  the  different  shape  of  the  opening  of  the  tem- 
poral fossa,  the  posterior  edge  of  which  is  much  wider  and  nearly  at  a 
right  angle  to  the  axis  of  the  skull.  If  therefore  a  line  is  drawn  across 
the  base  of  the  skull  at  the  level  of  this  edge,  the  line  passes  nearly 
through  the  palatine-basisphenoid  suture,  while  in  the  other  species  it 
crosses  the  basisphenoid  at  or  slightly  anterior  to  the  middle  (plates 
1,  fig.  1;  6,  30. 

The  skull  of  A.  mutica  is  unique  among  the  Amydas  in  the  slender- 
ness  and  delicacy  of  its  bones,  especially  those  of  the  snout,  which  is 
exceedingly  narrow  and  elongated.  The  distance  from  the  tip  to  the 
posterior  rim  of  the  orbit  is  much  greater  than  from  this  point  to  the 
posterior  edge  of  the  tympanic  cavity,  while  in  the  other  American 
species  it  is  much  shorter.  This  difference  is  mainly  caused  by  the 
reduction  in  A.  mutica  of  the  temporal  fossa,  the  diameter  of  which 
is  about  six  tenths  of  that  of  the  orbit,  while  in  the  other  species  it 
equals  or  averages  even  slightly  greater  than  the  orbit.  The  weakness 


SI  KJNEGER:    AMERICAN    SOFT-SHELL   TURTLES 


11 


of  the  A.  mutica  skull  is  further  emphasized  by  the  narrowness  of  the 
interorbital  spaee  and  the  practical  absence  of  an  alveolar  surface  on 
the  mandible. 

The  normal  skulls  of  the  remaining  American  species,  A.  ferox,  A. 
spinifera  and  A.  emoryi  are  essentially  alike  and  present  no  striking 
differences  from  the  normal  Amyda  skull.  They  agree  with  A.  agassizii 
in  the  greater  massiveness  of  the  bones  and  general  proportions  of  the 
parts.  With  A.  mutica  they  agree  in  the  longer  choanae  and  the  con- 
sequent shorter  distance  of  the  latter  from  the  intermaxillary  foramen. 


Table  1 


Cranial  measurements  reduced  to 
basicranial  length  of  skull 


(G 

c3 
fcC 

X 

o 

spinifera 

s- 
O 

s 

0) 

ferox  group 
collectively 

c3 

'-3 

< 

< 

< 

< 

< 

< 

Basicranial  length  in  millimeters, 

average 

51.8 

58.3 

53.8 

54.0 

42.8 

Range  of  b.  1.  of  specimens 

44-70 

45-70 

46-69 

44-69 

40-48 

Number  of  specimens  measured 

8 

12 

9 

9 

30 

6 

Tip  of  snout  to  orbit 

28.6 

24.9 

26.0 

25.2 

25.4 

28.71 

Orbit,  horizontal  diameter 

19.1 

20.0 

19.9 

20.2 

20.0 

20.6 

Orbit  to  tympanic  cavity 

31.7 

36.0 

27.1 

31.5 

31.5 

20.6 

Temporal  fossa,  longest  diameter 

19.3 

23.7 

19.3 

20.7 

21.2 

12.6 

Interorbital  width 

8.5 

6.0 

7.4 

7.8 

7.1 

5.1 

Maxillary  alveolar  surface,  width 

11.4 

6.8 

7.4 

8.9 

7.7 

5.1 

Internal  choanae,  length 

12.7 

15.4 

16.2 

15.4 

15.7 

14.7 

Choanae  to  intermaxillarv  foramen 

12.7 

7.7 

9.1 

8.9 

8.6 

9.8 

Intermaxillary  foramen,  length 

12.2 

11.3 

11.2 

9.8 

10.8 

15.6 

Mandibular  symphysis,  length 

19.5 

11.8 

13.2 

14.3 

13.1 

20.3 

Mandibular  alveolar  surface,  width 

10.6 

5.5 

5.4 

5.9 

5.6 

1  Four  specimens  only,  due  to  the  fact  that  the  bones — intermaxillaries  and  maxillaries  of 
this  species  are  so  fragile  or  poorly  ossified  that  they  were  lost  or  mutilated  in  the  preparation 
of  the  others.  The  omission  in  .4.  mutica  of  a  measurement  of  the  alveolar  width  of  the  mandible 
indicates  that  it  is  so  slightly  indicated  that  it  almost  may  be  said  to  be  nonexisting. 


12  bulletin:  museum  of  comparative  zoology 

The  great  changes  in  shape,  proportions,  color  pattern  and  structure 
of  the  soft -shelled  turtles  according  to  sex,  age,  and  individual  variabil- 
ity make  it  impracticable  to  construct  a  workable  key  to  the  species, 
hence  I  have  confined  myself  to  specimens  in  their  early  maturity, 
when  the  critical  characters  have  assumed  a  relative  and  comparable 
stability  unaffected  by  the  rapid  changes  of  youth  and  the  often  exag- 
gerated individual  peculiarities  of  old  age.  As  the  specific  differences 
in  the  skull  appear  fully  developed  and  most  easily  appreciated  in 
skulls  between  40  and  70  mm.  in  basicranial  length,  I  have  selected 
such  specimens  as  norms  to  which  other  sizes  may  be  more  or  less  suc- 
cessfully referred,  pointing  out  the  deviations,  as  shown  in  the  available 
material,  under  the  headings  of  the  various  species.  As  the  sexual 
differences  are  comparatively  slight  they  have  been  ignored  in  the  key, 
though  it  may  be  pointed  out  here  that  the  adult  males  differ  visibly 
from  the  females  in  the  tail  extending  considerably  beyond  the  cara- 
pace while  in  the  females  it  about  reaches  the  edge  of  the  disk;  in 
possessing  a  smaller  head  and  a  greater  expansion  of  the  plastral  cal- 
losities. 

It  is  finally  to  be  emphasized  that  specimens  may  be  met  with 
which  depart  so  far  from  the  norm  that  they  defy  positive  identifica- 
tion by  the  registered  characters  alone  or  in  combination.  Such  cases 
are  not  particularly  rare  among  the  Testudinata. 

Key  to  young  adult  specimens  of  North  American 
soft-shelled  turtles 

a1  Neurals  normally  8,  separating  all  the  pleurals;  entoplastron 
bent  at  an  angle  of  about  100°  or  more;  a  central  callosity  nor- 
mally on  entoplastron;  temporal  fossa  of  skull  small,  the  longest 
diameter  less  than  two  thirds  the  long  diameter  of  the  orbit; 
distance  from  orbit  to  tympanic  cavity  about  equals  diameter  of 
orbit;  intermaxillaries  (premaxillaries)  narrowly  touch  or  are 
separated  by  the  maxillaries.  Nostrils  rounded,  septum  between 
them  rather  wide  and  without  lateral  projecting  ridge;  no  tuber- 
cles on  the  leathery  disk  of  carapace  or  its  anterior  edge  {mulica 
group;  subgenus  Euamyda). 

Amyda  mutica,  (p.  14) 

a2  Neurals  normally  7  (occasionally  8),  last  pair  of  pleurals  in  con- 
tact; entoplastron  bent  at  an  angle  of  about  90°  or  less;  no  cal- 
losity on  entoplastron;  temporal  fossa  on  skull  about  equal  to 
or  longer  than  long  diameter  or  orbit;  distance  from  orbit  to 


stejneger:  American  soft-shell  turtles  13 

tympanic  cavity  much  greater  than  diameter  of  orbit;  maxil- 
laries  in  contact  above  intermaxillaries  (premaxillaries).  Nostrils 
crescent-shaped,  internal  ridge  projecting  on  each  side  from  the 
narrow  septum  between  them;  leathery  disk  of  carapace  with 
tubercles  at  least  on  anterior  edge. 

b1   Length  of  inner  bony  choanae  greater  than  their  distance 
from  intermaxillary  foramen;  mandibular  symphysis  less  or 
equal  to  length  of  choanae  {ferox  group). 
c1    wSculpture  of  bony  carapace  coarsely  grained,  usually  with 
numerous  irregular,  more  or  less  continuous  and  anasto- 
mosing longitudinal  ridges.    (Leathery  carapace  of  young 
with  longitudinal  rows  of  tubercles;  coloration  peculiar. 

Amyda  ferox,  (p.  25) 

c2    Sculpture  of  bony  carapace  finely  grained ;  (leathery  disk 
of  carapace  of  young  smooth ;  coloration  of  upper  surfaces 
gray,  more  or  less  marked  with  small  dusky  ocellae,  or 
solid  spots  or  lines;  plastron  uniform  white). 
d1   Tubercles  on  anterior  edge  of  leathery  carapace  well 
developed,  normally  triangular  and  pointed;  distinct 
tubercles  covering  anterior  and  posterior  flaps. 
e1    Bony   carapace    without   raised    bony    knobs    or 
"warts";  (young  with  one  marginal  dusky  line  on 
posterior  part  of  leathery  carapace). 

Amyda  spinifera,  (p.  43) 

e2  Adult  with  strong  tubercles  on  the  hind  part  of 
carapace  "supported  there  by  prominent  bony 
warts  of  the  bony  plates" ;  (in  young  margin  of  the 
leathery  carapace  marked  posteriorly  by  at  least 
two  parallel  dusky  lines). 

Amyda  spinifera  aspera,  (p.  56) 

d2   Tubercles  on  anterior  edge  of  leathery  carapace  poorly 

developed,  short  and  bluntly  rounded;  tubercles  on 

both  flaps,  if  present,  quite  minute,  except  in  very  old 

specimens. 

Amyda  emoryi,  (p.  65) 

b2  Length  of  inner  bony  choanae  equals  their  distance  from  in- 
termaxillary foramen;  mandibular  symphysis  longer  than 
length  of  choanae  (agassizii  group). 

Amyda  agassizii,  (p.  72) 


14  bulletin:  museum  of  comparative  zoology 


The  mutica  group 

The  only  species  so  far  known  in  this  group  deviates  in  its  characters 
and  their  combination  quantitatively  more  from  any  of  the  others  in 
the  genus,  so  much  in  fact  that  it  has  been  regarded  by  outstanding 
zoologists  such  as  Agassiz,  Cope,  True,  and  Baur  as  representing  a 
"good"  genus.  However,  the  differences  are  of  a  character  that  rather 
suggest  relationship  with  the  ferox  group  than  a  separate  phylogeny. 
The  difference  in  the  number  and  relations  of  neurals  does  not  seem  to 
have  any  genetic  significance  in  the  genus  Amyda.  The  long  drawn  out 
and  slender  snout  with  the  occasional  separation  of  the  maxillaries  at 
the  apex  and  the  shortening  of  the  temporal-tympanic  region  indicate 
modifications  due  to  some  food  specialization  with  which  the  absence 
of  the  dermal  ridge  in  the  nostrils  at  the  tip  of  the  proboscis  and  the 
thickening  of  the  septum  may  be  correlated.1  In  fact,  there  is  indica- 
tion of  the  ridge  inside,  though  not  reaching  the  opening  of  the  nostrils. 
The  extreme  development  of  the  plastral  callosities  is  purely  quan- 
titative and  is  closely  approached  in  old  specimens  of  Amyda  emoryi, 
and  a  small  callosity  on  the  entoplastron  is  often  observable  in  other 
specimens  of  the  genus.  The  more  circular  shape  of  the  body  outline 
is  rather  a  juvenile  character  with  which  the  obtuse  angle  of  the 
epiplastral  bones  is  correlated.  In  none  of  the  characters  does  the  A. 
mutica  show  any  approach  to  any  other  group  in  the  genus,  particu- 
larly not  to  any  of  the  Old  World  soft -shell  turtles,  so  that  there  seems 
to  be  no  convenience  in  recognizing  it  as  a  separate  genus.2 

Amyda  mutica3  (Lesueur) 
Plates  2,  3,  4 

1827.  Trionyx  muticus  LESUEUR,  Mem.  Mus.  Hist.  Nat.  Paris,  16,  Dec. 
1827,  p.  263,  pi.  7  (type-locality,  Wabash  River,  New  Harmony, 
Indiana;  type  Paris  Mus.  No.  787;  Lesueur,  collector ).— LE  CONTE, 
Ann.  Lye.  Nat.  Hist.  New  York,  3,  1830,  p.  95.— GRAY,  Syn.  Rept., 

1  Holbrook,  curiously  enough,  who  especially  called  attention  to  the  "two  characters  which 
always  exist",  viz.  the  "total  absence  of  spines  or  tubercles"  and  the  "great  difference  of  the 
nostrils"  in  describing  the  latter  speaks  of  them  as  "closely  approximated"  in  both  ferox  (in 
which  he  included  spinifera)  and  mutica  (North  American  Herpetology,  Edit.  2,  2,  p.  13  and 
p.  19)  while  in  reality  thev  "are  widely  apart"  in  the  latter,  as  pointed  out  bv  Agassiz  (Contrib. 
U.  S.  Nat.  Hist.,  1,  p.  398). 

'Should  the  statement  made  by  Dr.  Stockwell  (Journ.  Comp.  Med.  Surg.,  9,  1888,  p.  29) 
be  corroborated,  viz.,  that  "the  marginal  ossicles  in  A.  mutica  are  rudimentary;  in  A.  spinifer 
altogether  wanting,"  the  question  of  the  generic  status  of  A.  mutica  might  well  be  reopened. 

3  Latin,  muticus,  docked,  dehorned,  with  reference  to  the  absence  of  spines  on  the  anterior 
edge  of  the  leathery  carapace. 


stejxeger:  American  soft-shell  turtles  15 

pt.  1,  1831,  p.  46;  Cat.  Tort.  Brit.  Mus.,  1844,  p.  50;  Cat.  Shields 
Rept.  Brit.  Mus.,  p.  1,  March  8,  1856,  p.  69.— DUMERIL  AND 
BIBROX,  Erp.  Gen.,  2,  1835,  p.  482  (lapsus  for  Gymnopus).— HAR- 
LAN, Med.  Phys.  Res.,  1835,  p.  159  (Ohio  River  and  tributaries).— 
WIED,  Reise  Nord-Amerika,  1,  pt.  3,  1838,  p.  140  (Pittsburgh,  Pa.). 
-HOLBROOK,  North  Amer.  Herpet.,  1  ed.,  4,  1840,  p.  17,  pi.  2 
(Mississippi  and  tributaries);  2  ed.,  2,  1842,  p.  19,  pi.  2  (Mississippi 
and  tributary  streams).— DE  KAY,  Zool.  New  York,  Rept.,  1842, 
p.  7  (Ohio  River).— TROOST,  Seventh  Geol.  Rep.  Tennessee,  1844, 
p.  39  (Tennessee;  milieus  misprint). — DUMERIL,  Cat.  Meth. 
Rept.  Mus.  Paris,  pt.  1,  1851,  p.  22  (types).— STRAUCH,  Mem. 
Acad.  Sci.  St,  Petersbourg,  ser.  7,  5,  No.  7,  1862,  p.  174;  8,  No.  13, 
1865,  p.  125;  38,  No.  2,  1890,  p.  118.— HOY,  Geology  of  Wisconsin, 
1,  1883,  (p.  423)  (Wisconsin).— BOULENGER,  Cat.  Chel.  Brit. 
Mus.,  1889,  p.  260,  fig.  68  (Mississippi,  Ohio  and  Saint  Lawrence). — 
HAY,  Indiana  Geol.  17  Rep.,  1892,  p.  551;  Batr.  Rept.  Indiana, 
1893,  p.  143  (Indiana:  Delphi;  Madison;  Terre  Haute.  Illinois: 
Mt.  Carmel).— HURTER,  Trans.  Acad.  Sci.  St.  Louis,  6,  1892,  p. 
259  (Missouri:  Mississippi  River  near  St.  Louis).— SIEBENROCK, 
Sitz.  Ber.  Akad.  Wiss.  Wien,  Math.-Nat.  KL,  111,  1902,  p.  822,  fig.  5 
(plastron);  Zool.  Jahrb.  Suppl.,  10,  pt.  3,  1909,  p.  605;  Ann.  Naturh. 
Hofmus.  Wien,  27,  1913,  p.  214,  sep.  p.  44  (plastron);  Verh.  Zool. 
Bot.  Ges.  Wien,  73,  Aug.  1923,  p.  192.— DITMARS,  Reptile  Book, 
1907,  p.  78,  pi.  27,  low.  fig.  (St.  Louis,  Mo.).— OVER,  South  Dakota 
Geol.  Nat.  Hist.  Surv.  Bull.  12,  Oct.  1923,  p.  18,  pi.  7  (Missouri  Riv. 
and  eastward,  South  Dakota). 

Gymnopodus  muticus  DUMERIL,  Arch.  Mus.  Hist.  Nat.  Paris,  7, 
1855,  p.  203. 

Gymnopus  muticus  DUMERIL  and  BIBRON,  Erpet,  Gen.  9,  1854, 
p.  236.— WIED,  Nova  Acta  Leopold.-Carol.,  32,  pt.  1,  1865,  p.  54 
(Ft.  Mackenzie,  Missouri  River,  6-8  miles  below  Cedar  Isl.,  South 
Dakota). 

Amyda  mutica  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  1,  1857,  p. 
399;  vol.  2,  pi.  6,  figs.  6-7  (Lake  Erie  and  Ontario;  Delphi,  Ind.; 
Burlington,  Iowa;  Osage  River,  Missouri;  Alleghany  Riv.,  Pa.). — 
GRAY,  Suppl.  Cat.  Shield  Rept.  Brit.  Mus.,  p.  1,  1870,  p.  95.— 
COPE,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  1875,  p.  41  (middle  and  northern 
tributaries  of  Mississippi  and  the  St.  Lawrence  Riv.). — JORDAN, 
Man.  Vert.  North.  United  States,  ed.  3,  1880,  (p.  168);  ed.  8,  1899, 
p.  206  (Canada  to  Ohio  River,  and  N.  W.).— CRAGIN,  Trans.  Kan- 
sas Acad.  Sci.,  7,  1881,  p.  116  (Kansas:  Manhattan;  Blue  and  Kansas 
Rivers).— SMITH,  Rep.  Geol.  Surv.  Ohio,  4,  1882,  p.  668  (Ohio 
River,  Ohio).— TRUE,  Bull.  U.  S.  Nat.  Mus.,  No.  24,  1883,  p.  28 
(Madison,  Ind.;  Mt.  Carmel,  111.;  St.  Louis,  Mo.;  Ft.  Smith,  Ark.); 
Fish  Industr.  United  States,  sect.   1,  1884,  p.   152.— HOY,  Geol. 


16  bulletin:  museum  of  comparative  zoology 

Wisconsin  Survey  1873-1879,  1,  1883,  p.  423  (tributaries  to  the  Mis- 
sissippi in  Wisconsin). — DAVIS  and  RICE,  Illinois  State  Lab.  Nat. 
Hist.  Bull.  No.  5,  1883,  p.  52  (North  of  Ohio  River);  Bull.  Chicago 
Acad.  Sci.,  1,  No.  3,  1883,  p.  32  (Illinois).— BAUR,  Zool.  Anz.,  10, 
1887,  p.  99  (descr.  plastron) ;  Amer.  Natural.,  22,  1888  (1122);  Proc. 
Amer.   Philos.   Soc,   31,   1893,    (p.   220).— STOCKWELL,   Journ. 
Comp.  Med.  Surg.,  9,  No.  1,  Jan.  1888,  p.  29.— HIGBY,  Trans. 
Wisconsin  Acad.  Sci.,  7,  1889,  p.  159  (Wisconsin:  western  half  of 
state).— GARM AN,  H.,  Bull.  Illinois  Lab.  Nat.  Hist.,  3,  1892,  p. 
247  (throughout  Illinois:  Mackinaw  Creek,  Woodford  Co.;  Quincy; 
Illinois  Riv.,  Peoria;  Wabash  Riv.,  Mt.  Carmel;  Ohio  Riv.,  Cairo). — 
RHOADS,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  1895,  p.  404  (Ten- 
nessee).— GAGE,  Trans.  American  Microsc.  Soc,  17,  1895,  (p.  185) 
(embryology).— SMITH,  Proc.  Linn.  Soc.  New  York,  1898-1899, 
No.  11  (1899)  p.  24  (not  near  New  York  City).^-McLAIN,  Notes 
Coll.  Rept.  Arkansas,  1899,  p.  1  (Ft.  Smith,  Ark.).— ATKINSON, 
Ann.  Carnegie  Mus.  Pittsburgh,  1,  1901,  p.  154  (Allegheny  Co., 
Pa.).— PAULMIER,  Bull.  51,  New  York  State  Mus.,  1902,  p.  392 
(probably  in  northern  New  York  State). — MORSE,   Proc.   Ohio 
Acad.  Sci.,  4,  pt.  3,  Spec.  Pap.  No.  9,  1904,  p.  138  ("not  recorded  for 
Ohio")- — ?NASH,  check  list  of  Vertebrates  of  Ontario,  Batrachians, 
Reptiles,     Mammals,     1906,    p.    17    (Ontario,    Lake    Erie,    rare). 
SURFACE,    Zool.    Bull.    Pennsylvania    Dep.    Agric,  6,  nos.  4-5, 
Sept.  1,  1908,  p.  119  (Pennsylvania,  liable  to  be  found  in  Erie 
County  and  tributaries  of  the  Ohio).  HURTER  and  STRECKER, 
Trans.    Acad.    Sci.  St.  Louis,   18,   no.  2,   1909,   p.   21    (Arkansas: 
Ft.   Smith;   Pink   Bluff;  Little  Rock).  —  HURTER,  Herpet.   Mis- 
souri, 1911,  p.  249   (Mississippi,   Osage,   Gasconade  and  Meranac 
Rivers).— CLARK   and   SOUTHALL,   Rep.  U.    S.    Comm.  Fish., 
1919     (1920)     App.     No.     7,     p.    15    (economics).  —  MULLER, 
Amer.    Midland    Natural.,    7,    no.   6,   Nov.   1921,  p.   180    (Iowa: 
Fairport;     habits).  —  BLANCH ARD,     Occas.     Pap.    Zool.    Mus. 
Univ.  Michigan,  No.  117,  July  6,  1922,  p.  18  (Trotter's  Landing, 
Benton  Co.,  Tenn.).— WEED,  Copeia,  No.  116,  March  15,  1923,  p. 
48  (Meredosia,  Ills.).— PRATT,  Man.  Vert.  United  States,  1923,  p. 
249.— STRECKER,  Baylor  Univ.  Bull.,  27,  No.  3,  Sept.  1924,  p.  47 
(Little  Rock,  Ark.);  Contr.  Baylor  Univ.  Mus.,  No.  7,  July  15,  1926, 
p.  7  (Neches  River,  Henderson  Co.,  Texas,  probably). — MYERS, 
Proc.  Indiana  Acad.  Sci.,  35,  1926,  p.  292  (Indiana).— STRECKER 
and  FRIERSON,  Contr.  Baylor  Univ.  Mus.,  No.  5,  May  15,  1926, 
p.    10   (Caddo  and  De  Soto  Parishes,   La.).— ORTENBURGER, 
Proc.  Oklahoma  Acad.  Sci.,' 6,  pt.  1,  1926,  p.  100  (McCurtain  and 
Pushmataha   Cos.,    Oklahoma).— STRECKER   and    WILLIAMS, 
Contr.  Baylor  Univ.  Mus.,  No.  12,  Dec.  27,  1927,  p.  16  (Christoval, 
Tom  Green  Co.,  Texas);  No.  17,  Oct.  20,  1928,  p.  19  (Bowie  Co.:  "no 


stejneger:  American  soft-shell  turtles  17 

doubt")-— BURT,  Occas.  Pap.  Mus.  Zool.  Univ.  Michigan,  No.  189, 
Dec.  12,  1927,  p.  9  (Manhattan,  Riley  Co.,  Kansas) ;  Amer.  Midland 
Natural.,  16,  No.  3,  1935,  p.  321  (Barber  and  Reno  Cos.,  Kansas).— 
POPE  and  DICKINSON,  Bull.  Publ.  Mus.  Milwaukee,  8,  No.  1, 
April  3,  1928,  p.  82,  pi.  21,  figs.  5-6  (Wisconsin:  Mississippi  River 
counties;  Crawford  and  Pepin  Cos.). — JORDAN,  Man.  Vert. 
Northeast,  U.  S.  (13  ed.)  1929,  p.  254  (Middle  and  Northern  tribu- 
taries, Miss,  and  St.  Lawrence  rivers.) — ORTENBURGER  and 
FREEMAN,  Publ.  Univ.  Oklahoma  Biol.  Surv.,  2,  1930,  p.  188 
(Oklahoma:  Alfalfa  and  Comanche  Cos.). — WALKER,  Copeia, 
1931,  No.  1,  p.  12  (Scioto  and  Brown  Cos.,  Ohio).— BOYER  and 
HEINZE,  Trans.  Acad.  Sci.  St.  Louis,  26,  No.  4,  Apr.  1,  1934,  p.  199 
(Missouri:  Jefferson  Co.:  Meranac  River). — RUST,  Blatt.  Aquar. 
Terrarienk.,  45,  1934,  p.  ;  sep.  p.  12.— TAYLOR,  Univ.  Kansas 
Sci.  Bull.,  22,  No.  11,  Apr.  15,  1935,  p.  218  (Arkansas:  Duvall  Bluff, 
Prairie  Co.;  Lewisville,  Lafayette  Co.).— DELLINGER  and 
BLACK,  Occas.  Pap.  Univ.  Arkansas,  No.  1,  June  1938,  p.  46  (Ar- 
kansas: Garland,  Jefferson,  Lafayette,  Prairie,  Pulaski,  Sebastian, 
Franklin  and  Lawrence  Cos.). — SOLA,  Bull.  New  York  Zool.  Soc, 
34,  No.  5,  Sept.-Oct.,  1931,  pp.  134,  142,  155,  fig.  7  upper  (western) 
part  of  Pennsylvania  along  Lake  Erie). — CAHN,  Illinois  Biol. 
Monogr.,  16,  Nos.  1-2,  Aug.  31,  1937,  p.  176;  pis.  24,  30  fig.  a;  map 
19(Illinois).— CONANT,  Amer.  Midland  Natural.,  20,  No.  1,  July, 
1938,  p.  154,  pi.  21,  fig.  1  (left),  fig.  2  (Ohio:  Scioto,  Muskingum  and 
Ohio  Rivers;  map).— PARKER,  Rep.  Reelfoot  Lake  Biol.  Sta.,  3, 
Jan.  1939,  p.  88  (Tennessee:  Reelfoot  Lake,  nowhere  abundant). — 
WELTER  and  CARR,  Copeia,  1939,  No.  3,  Sept.  9,  p.  130  (Ken- 
tucky, east.:  Triplet  Co.;  Fox,  Fleming  Co.;  rare). — LOGIER,  Roy. 
Ontario  Mus.  Zool.  Hanb.  No.  4,  p.  57  (Ontario:  Lake  Erie,  probably 
misidentified).— GENTRY,  Rep.  Reelfoot  Lake  Biol.  Sta.,  5,  Jan. 
1941,  p.  75  (Tennessee:  Pickett  Co.);  Journ.  Tennessee  Acad.  Sci., 
16,  No.  3,  1941,  p.  332  (Tennessee:  Obey  River,  Pickett  Co.).— 
ANDERSON,  Bull.  Chicago  Acad.  Sci.,  6,  No.  11,  1942,  p.  219 
(Missouri:  Jackson  Co.:  Fry's  Lake).— PETERS,  Copeia,  1942,  No. 
3,  Oct.  8,  p.  183  (Illinois:  Cumberland  Co.). 
1864.  Potamochdys  ?  microcephala  GRAY,  Proc.  Zool.  Soc.  London,  1864 
(p.  87)  (type-locality,  Sarawak,  Borneo!!;  type  in  British  Museum). 
Callinia  microcephala  GRAY,  Proc.  Zool.  Soc.  London,  1869  (p. 
222);  1873,  p.  62,  fig.  11;  Suppl.  Cat.  Shield  Rept.  Brit.  Mus.,  pt.  1, 
1870,  p.  108;  Hand-list  Shield  Rept.  Brit.  Mus.,  1873,  p.  83. 

Types.     In  the  Musee  d'Histoire  Naturelle  at  Paris,  No.    787   is 
a  shell  on  the  plastron  of  which  is  written:  "Trionix  mutica  Lesueur. 

Wabash  River  par  moi aout  1S27",  and  on  a  paper  label  pasted 

on  the  underside  of  the  stand:  "Tortue  qui  n'a  point  le  bord  desou 


18  bulletin:  museum  of  comparative  zoology 

disque  spineaux  &  que  j'ai  designee  sou  le  nom  de  mutica  dans 

precedente  note  accompagniee  de  figure  que  je  vous  ai  fait  passer  |?] 
C.  A.  Lesueur."  It  is  designated  as  the  type  on  the  printed  label.  The 
bony  carapace  is  106  mm.  in  length  and  107  mm.  in  width.  A  pair  of 
fontanelles  between  first  pair  of  pleurals ;  8  neurals  separate  the  pleurals. 

No.  788  is  a  cotype  retained  by  Lesueur  until  1844  when  it  was  ac- 
quired by  the  Museum.  It  is  a  larger,  mounted  specimen,  the  bony 
carapace  measuring  134  mm.  in  length,  147  mm.  in  width;  fontanelles 
obliterated;  8  neurals,  but  last  pair  of  pleurals  broadly  in  contact. 
Underside  of  stand  inscribed  "Gymnopus  muticus  Lesueur.  Wabash. 
Acquis  de  Mr.  Lesueur  1844"  (Lesueur  died  December  12,  1846). 

The  so-called  "Spineless"  or  "Brown  Softshell  Turtle",  the  smaller 
of  our  American  species,  may  be  easily  recognized  by  the  characters 
given  in  the  key.  In  the  adolescent  and  adult  carapaces  the  absolute 
absence  of  tubercles  on  the  anterior  edge  of  the  leathery  disk  will 
identify  the  A.  mutica  even  when  the  head  is  missing  or  the  proboscis 
is  so  mutilated  as  to  defy  examination.  At  the  stage  when  the  very 
young  specimens  of  all  the  species  are  nearly  circular  in  outline  and  the 
tubercles  on  the  anterior  edge  of  carapace  of  those  species  normally 
possessing  them  may  in  some  individuals  be  so  indistinct  as  to  be 
doubtful,  the  oval  nostrils  without  the  septal  tubercle  will  positively 
identify  A.  mutica. 

The  other  differential  characters  keyed  are  not  always  to  be  relied 
on  because  of  individual  variation.  Thus  while  normally  the  number  of 
pleurals  is  7  pairs,  separated  the  entire  length  of  the  bony  carapace  by 
a  series  of  8  neurals,  there  are  many  and  significant  exceptions.  Thus, 
as  already  mentioned,  the  mounted  cotype  of  the  species  in  the  Paris 
Museum  has  the  last  pair  of  pleurals  broadly  in  contact.  Similarly  the 
U.S.N.M.  102910  has  the  eighth  pair  of  pleurals  in  contact  for  at  least 
half  their  length;  No.  54734  has  all  the  pleurals  separated  by  the  8 
neurals,  but  it  has  only  6  pleurals  on  the  right  side,  against  7  on  the 
left;  U.S.N.M.  92605,  95134  and  029261  have  only  7  neurals  and  7 
pairs  of  pleurals  and  the  last  pair  is  broadly  in  contact  behind  the 
neurals.  While  visiting  the  Museum  of  Comparative  Zoology  during 
the  early  days  of  these  investigations,  I  examined  two  specimens,  both 
unfortunately  at  that  time  without  numbers  and  locality,  one  adoles- 
cent with  the  seventh  pair  of  pleurals  in  contact  behind  the  neurals, 
while  the  second,  an  adult  skeleton  with  a  bony  carapace  measuring 
140  mm.  in  length  and  150  mm.  in  width,  was  still  more  abnormal 
having  8  pleurals  on  the  right  side,  against  7  on  the  left,  and  with  9 
neurals  separating  all  the  pleurals. 


stejneger:  American  soft-shell  turtles  19 

The  greater  angular  width  of  the  entoplastron  is  quite  characteristic 
of  this  species,  but  it  is  sometimes  questionable  in  application  because 
of  difficulty  of  exact  measurement.  However,  it  is  useful  where  other 
characters  are  irregular  or  in  case  one  has  to  identify  a  disassociated 
plastron  or  a  single  bone.  The  great  extension  of  the  plastral  callosities 
in  the  males  is  also  a  character  of  value,  though  the  callosities  on  old 
male  A.  emoryi  may  reach  similar  proportions  on  the  hyo-,  hypo-,  and 
xiphi-plastra.  A  central  callosity  is  normally  present  in  A.  mutica  on 
the  entoplastron  though  exceptional  in  the  other  species.  Small  callosi- 
ties on  the  epiplastra  are  not  uncommon  in  A.  midica,  though  rare  in 
the  others. 

The  sutural  meeting  of  the  maxillaries  on  the  upper  side  of  the 
snout  above  the  premaxillary  (intermaxillary)  is  one  of  the  characters 
of  the  trionychid  skulls,  and  is  probably  a  normal  or  at  least  original 
condition  in  A.  mutica,  but  as  noted  above  the  extreme  tapering  and 
hence  weakening  of  the  snout  in  this  species  results  in  the  frequent 
loss  of  these  parts  in  the  preparation  of  the  skulls.  In  my  series  of 
A.  mutica  skulls  there  are  only  four  perfect  specimens  and  in  one  of 
these,  No.  102677,  the  maxillaries  are  plainly  in  contact  on  the  upper 
side,  above  the  premaxillary,  while  in  the  others,  Nos.  53521,  54733  and 
029261,  the  maxillaries  are  separated  by  the  premaxillary. 

A  negative  character  of  the  young  A.  mutica  is  the  absence  of 
ocellated  or  solid  black  rounded  spots  on  the  carapace,  and  of  a  defined 
angular  figure  on  top  of  the  snout  at  the  base  of  the  proboscis.  In  a  gen- 
eral way  the  coloration  is  less  distinctive  than  in  the  other  species.  The 
yellowish  margin  of  the  leathery  carapace  seems  to  be  definitely  wider. 

The  coloration  and  pattern  in  this  as  well  as  the  other  species  of 
the  genus  becomes  gradually  more  obscure  with  age,  and  varies  in- 
dividually as  well  as  locally  according  to  environmental  conditions. 
It  may  therefore  not  be  unwelcome  if  I  include  a  few  color  descriptions 
of  living  or  freshly  killed  specimens  which  have  come  under  my  ob- 
servation when  comparison  with  Ridgway's  "Nomenclature  of  Colors 
for  Naturalists"  (1886)  was  possible. 

On  September  4,  1934,  the  National  Museum  received  from  C.  R. 
Rogers  two  live  specimens  taken  in  Medicine  Lodge  River,  one  mile 
SW  of  Lake  City,  Barber  Co.,  Kansas.  Description  was  at  once  made: 
U.S.N.M.  No.  95185,  young  adult  female  (leathery  disk  about  200 
mm.).  Iris  bright  "buff",  the  ring  nicked  slightly  in  front  and  behind 
by  a  small  blackish  spot. — General  color  above  nearly  uniform 
"tawny"  with  very  faint  mottlings  of  lighter  "tawny-ochraceous"  and 
darker  "raw  umber",  especially  on  the  posterior  flap,  on  which  a  faint 


20  bulletin:  museum  of  comparative  zoology 

submarginal  dusky  line  borders  the  pale  margin  which  is  lightly 
suffused  with  "rufous";  top  and  sides  of  head  like  back  with  a  sharply 
defined  band  of  "ochraceous  buff"  narrowly  edged  with  dusky;  the 
band  continues — though  fainter — anteriorly  through  the  eye  on  to 
the  canthus  rostralis  converging  towards  the  base  of  the  proboscis 
without  meeting  that  of  the  other  side;  from  the  side  of  the  occiput 
indication  of  the  band — though  more  irregular — on  the  side  of  the 
neck;  underside  with  a  fine  network  of  red  blood  vessels  shining 
through  imparting  a  pinkish  tinge  to  the  soft  parts  which  fades 
gradually  through  "lavender"  and  "pearl  blue"  into  "china-blue"  on 
the  palms,  soles  and  digits  merging  on  the  underside  of  the  webs  into 
"tawny  ochraceous"  exteriorly  and  more  pinkish  interiorly;  throat  and 
chin  like  the  soles;  lower  lips  whitish;  callosities  pale  "fawn-color" 
tinged  centrally  with  blueish. 

The  other  specimen,  No.  95186,  is  a  much  smaller  male  (leathery 
carapace  length  about  135  mm.)  (pi.  3).  Iris,  a  pale  yellowish  ring,  but 
the  black  spots  are  somewhat  larger  than  in  the  older  specimen. 
Colors  are  also  essentially  like  the  latter,  but  the  "ochraceous  buff" 
postocular  band  has  the  edges  even  better  defined  and  on  the  side  of 
the  neck  joins  the  pale  color  of  the  underside  which  is  sharply  set  off 
from  that  of  the  upper  side  and  extends  onto  the  upper  lip;  the  band 
is  only  indicated  in  front  of  the  eye  by  a  small  elongated  triangular 
spot  of  pale  "ochraceous". 

About  the  same  time  the  National  Museum  received  two  live  speci- 
mens from  J.  H.  Hall,  collected  in  Mississippi  (Marion  County,  Colum- 
bia), both  females.  No.  95133,  the  larger  one  (disk  approximately  185 
mm.  long).  Iris  bright  buff  yellow  with  a  black  horizontal  bar.  General 
color  above  "clay  color"  with  irregular  blotches  of  pale  "raw  sienna"; 
marginal  dark  ring  on  carapace  broken,  faint,  "hair-brown";  upper 
side  of  neck  washed  with  "tawny  ochraceous";  postocular  stripe  dull 
"buff -yellow",  edged  with  dusky  ("hair-brown");  front  legs  above  as 
well  as  dorsal  and  lateral  surface  of  neck  sprinkled  with  small  dusky 
spots;  hind  feet  pale  "olive  yellow"  with  slightly  larger  and  darker 
dots  and  marblings;  webs  verging  on  "clay  color";  underside  of  plas- 
tron "flesh  color",  which  on  the  white  ground  of  the  legs  changes  into 
"pale  blue"  and  on  the  soles  and  front  feet  verges  on  "heliotrope 
purple";  underside  of  webs  pinkish  towards  the  edge;  claws  horny 
white;  callosities  (none  on  epiplastra)  "vinaceous-cinnamon".  No  fork 
figure  on  top  of  snout;  no  spots  or  definite  dusky  markings  on  soles. 

The  smaller  specimen,  No.  95134  (leathery  disk  approximately  155 
mm.)  essentially  as  the  larger  one. 


stejneger:  American  soft-shell  turtles 


21 


Table  2 
Cranial  measurements  of  mutica  in  millimeters 


CD 
M 

03 

0J 

M 

03 

O 

a 

of 
PE 

O 

I— 1 

l-c 

O 

a 

o3 

O? 

& 

O 

e*- 

i 

a 

o3 

M 

+3 

O 

a 
'3 

oT 

o 
1— 1 

O 

o 
"a; 

aT 

03 
CO 

a> 

5 

H 

43 

o 

o 

53 
« 

aT 

CD 

co 

CO 
0) 

a 
a 

CD 

H 

co 

C 

CD 

e 

0J 

ft 

CO 

o 

CD 
bfl 

<M 
iO 

co 

41.0 
11.0 

i— i 

CO 
CM 
OS 

cn 

o 

42.0 
12 

00 

i— i 

CO 

43.0 

co 

<N 

O 

48.0 
14.5 

o 

OS 
(N 

O 

T— t 

CO 

> 

Basicranial  length 

40.0 
11.5 

42.8 

Tip  of  snout  to  orbit 

12.3 

Orbit,  horizontal  diameter 

8.0 

8.5 

p 

9.0 

9.0 

9.5 

7.0 

8.6 

Orbit  to  tympanic  cavity 

8.0 

8.0 

8.5 

9.0 

10.0 

9.0 

8.8 

Temporal  fossa,  longest  di- 

ameter 

5.0 

4.-r 

5 

6.0 

5.0 

6.0 

5.0 

5.3 

Interorbital  width 

2.0 

2.0 

..0 

2.0 

2.0 

3.0 

2.0 

2.1 

Maxillary  alveolar  surface, 

width 

2.0 

2.5 

2.0 

1.5 

2.0 

2.5 

2.0 

2.1 

Internal  choanae,  length 

6.0 

7.0 

6.5 

6.0 

6.0 

6.5 

7.0 

6.7 

Choanae  to  intermaxillary 

foramen 

4.0 

4.0 

4.0 

4.0 

4.5 

5.0 

4.0 

4.2 

Intermaxillary  foramen, 

length 

6.0 

6.5 

6.5 

8.0 

6.5 

6.7 

Mandibular  symphysis, 

length 

8.0 

8.5 

8.0 

9.0 

10.0 

8.0 

8.6 

Mandibular  alveolar  sur- 

face, width 

The  colors  of  a  freshly  killed  male  specimen  (leathery  disk  204  mm. 
long)  collected  by  Dr.  C.  E.  Burt  in  Kansas,  Reno  County,  6  miles  E. 
of  Turon  (U.S.N.M.  No.  95259)  were  as  follows:  Ground  color  "broc- 
coli brown"  mottled  with  numerous  very  irregular  and  more  or  less 
anastomizing,  ragged-edged  "sepia"  spots  occupying  as  much  space  as 


22  bulletin:  museum  of  comparative  zoology 

the  ground  color;  posterior  edge  of  disk  pale  tinged  with  "burnt  um- 
ber"; no  submarginal  blackish  line;  upper  soft  parts  same  broccoli 
brown  with  very  small  and  faint  irregular  "sepia"  dots;  a  very  pale 
"russet"  band,  raggedly  edged  with  blackish,  from  canthus  rostralis 
through  eye  and  over  ear  slanting  on  side  of  neck  halfway  down  the 
neck ;  pale  lines  in  front  of  eyes  do  not  meet  those  on  snout  nor  is  there 
trace  of  a  connecting  line  forming  fork  or  triangle  at  base  of  proboscis ; 
underside  whitish  with  a  faint  glaucous  blue  gray  tinge  on  throat, 
underside  of  neck  and  feet;  no  dusky  markings  on  feet;  callosities,  in- 
cluding the  small  central  one  on  entoplastron,  pale  blueish  "plumb- 
eous". 

Geographical  Distribution 

Mississippi  River  and  tributaries;  north  to  South  Dakota  and 
Minnesota;  east  to  western  Pennsylvania;  west  to  Kansas,  Oklahoma 
&  Texas. 

Recorded  from  northern  localities  on  the  Trinity,  Brazos  and 
Colorado  Rivers. 

Agassiz  (Contr.  Nat.  Hist.  U.  S.,  1,  p.  404,  footnote)  writes,  "De- 
Kay's  Trionyx  ocellatus  is  Amyda  mutica,"  a  statement  which  seems 
to  be  erroneous.  The  reference  to  DeKay's  Trionyx  ocellatus  appears  to 
be  the  following  note  in  his  Zoology  of  New  York  (pt.  3,  1842,  p.  7) 
under  Trionyx  ferox  [DeKay=  T.  spiniferus]:  "The  description  given 
above  [p.  6:  "anterior  margin  in  the  adult  with  numerous  pointed 
tubercles,  which  may  be  faintly  and  distantly  traced  in  the  young"] 
was  taken  several  years  since,  from  a  specimen  obtained  in  the  Mohawk 
River.  .  .  .  The  specimen,  as  I  then  thought,  varied  so  much  from  any 
description  of  the  ferox  within  my  reach,  that  I  considered  it  to  be  new, 
and  named  it  ocellatus."  The  description  of  the  Mohawk  River  speci- 
men is  clearly  that  of  an  adult  A.  spin  if  era,  and  cannot  be  taken  as  the 
record  of  A.  mutica  in  the  Mohawk  River.  DeKay's  mention  that  he 
suspected  ferox  and  muticus  to  be  identical  probably  caused  Agassiz's 
statement. 

List  of  specimens  in  U.  S.  National  Museum 


4783  (2)  <?  a< 

id.,  d"  adol.         ? 

? 

7646  d"  adol. 

Mo.,  St.  Louis 

Engelmann 

7647  juv. 

U              tl 

? 

7655  juv. 

Miss.,  Monticello 

Helen  Tennison 

7659-60  (2) 

111. 

Kennicott 

7746 

Ark.,  Arkansas  R.  near 

Ft.  Smith 

Lt.  Whipple 

stejneger:  American  soft-shell  turtles 


23 


8337  9 

ad. 

Indiana,  Madison 

9615  c? 

adol. 

111.,  Mt.  Carmel 

Mrs.  M.  E.  Turner 

9646  d1 

adol. 

11                     tt 

Robert  Ridgway 

May 

1878 

9647  d"  adol. 

II                  tt 

tt 

(« 

9650  <?  adol. 

it                   a 

tt 

a 

9651  rf"  adol. 

t;                  it 

tt 

tt 

9727  9 

ad. 

it                  ll 

J.  Schneck 

n 

11629  juv. 

? 

? 

11630  9 

adol. 

? 

? 

12794  (2. 

I  juv. 

< 

?& 

111.,  Mt.  Carmel 

J.  Schneck 

13549 

? 

? 

14780  juv. 

Mo.,  St.  Louis 

J.  Hurter 

Dec. 

20,  1887 

19626  9 

ad. 

? 

? 

19627  9 

adol. 

o 

? 

21418  9 

? 

? 

22629  juv. 

Tex.,  Sabine  R.,  5  mi.  S. 

Jordan  &  Gilbert 

1884 

Longview. 

029261  ad 

. 

? 

Dr.  G.  Baur 

45735-8 

Iowa,  Fairport,  Missis- 

sippi R. 

Snyder 

June 

2,  1916 

52528  <?  ad. 

Kansas 

Roy  L.  Moodie 

52116-8 

juv. 

111.,  Olney 

Robert  Ridgway 

53521    9 

ad. 

Iowa,  Fairport 

Bur.  Fisheries 

Apr. 

24,  1911 

54733  9 

ad. 

ii             it 

J.  Snyder 

54734   9 

ad. 

it            tt 

tt 

Aug. 

8,  1916 

54742 

"     (Missis- 

sippi R.) 

a 

May 

8,  1916 

55525 

Mo.,  St.  Louis 

J.  Hurter 

Mar.  23,  1907 

55526 

it 

a 

May 

14,  1913 

55527 

Ark.,  Jeff.  Co. 

tt 

1899 

55528 

"      Little  Rock 

tt 

June 

1,  1900 

55600 

Tex.,  McLennan  Co. 

J.  Hurter 

1896 

59267  9 

ad. 

Mo.,  Alexandria 

E.  Stringham 

June 

5,  1916 

59268-9 

Minn.,  Homer 

F.  Schrader 

Sept. 

8,  1916 

59276 

111.,  betw.  Warsaw  & 

Hamilton 

E.  Stringham 

Aug. 

23,  1916 

59278  9 

ad. 

Mo.,  Alexandria 

June 

5,  1916 

59281 

Iowa,  Keokuk 

Aug. 

16,  1916 

59282 

tt           it 

June 

7,  1916 

59283 

it           it 

July 

1,  1916 

59284 

it           it 

June 

7,  1916 

59982 

"Central  U.S." 

O.  P.  Hay 

60054-6 

Iowa,  Fairport 

Bur.  Fish. 

60561 

Mo.  St.  Louis 

J.  Hurter 

24 


bulletin:  museum  of  comparative  zoology 


71547 

Okla.,  6  mi.  E.  Ingersoll 

A.  I.  Ortenburger 

92605  cf  adol. 

Miss.,  Greenville 

S.  F.  Hildebrand 

May  29,  1933 

95133-4  9  adol. Miss.,  Columbia 

J.  H.  Hall 

Aug.  1934 

95185  9  ad. 

Kans.,  1  mi.  S.W.  Lake 

City 

C.  R.  Rogers 

Aug.  31,  1934 

95186  d1  juv. 

Kans.,  1  mi.  S.W.  Lake 

City 

<( 

it 

95259  a* 

Kans.,  6  mi.  E.  Turon 

C.  E.  Burt 

May  25,  1934 

95260  9  adol. 

U                      U                       {{ 

it 

it 

95415   9  adol. 

"       3  mi.  S.E.  Oxford 

u 

1934 

100422   9  adol. 

La.,  Rayville 

it 

Aug.,  1935 

100813  pull. 

Kans.,  Wakeeney 

O.  P.  Hay 

102612-3 

111.,  5  mi.  S.  Savannah 

P.  Bartsch 

July   29,  1907 

102677   9  ad. 

Tenn.,  Reelfoot  Lake 

W.  M.  Perrygo  & 

C.  Lingebach 

May     6,  1937 

102910   9  ad. 

it                     it               a 

W.  M.  Perrygo  & 

C.  Lingebach 

May     8,  1937 

115939 

[Miss.] 

[B.  C.  Wailes] 

The  ferox  Group 

Although  quite  distinct,  the  three  species  included  form  a  rather 
close  group  chiefly  characterized  by  the  uniformity  of  their  skulls. 
Common  for  all  three  is  the  large  opening  of  the  inner  choanae  with  the 
concomitant  shortness  of  the  distance  of  the  latter  from  the  inter- 
maxillary foramen,  which  distinguish  them  from  the  agassizii  group, 
while  the  normal  proportion  of  the  preorbital  region  sets  them  off  from 
the  mutica  group.  This  statement  may  seem  strange  in  view  of  the 
opinion  of  many  early  competent  herpetologists  who  refer  to  the  species 
of  the  ferox  group  under  two  different  generic  terms,  a  situation  caused 
by  the  confusion  of  the  identity  of  the  specimens  which  served  as  basis 
for  the  generic  concept,  as  will  be  shown  later  on. 

Externally  the  ferox  group  differs  from  the  mutica  group  by  the 
crescentic  shape  of  the  nasal  openings,  a  character  shared  by  all  the 
other  species  of  the  genus,  and  by  the  presence  of  the  dermal  tubercles 
on  the  carapace.  From  the  agassizii  group,  however,  there  is  no  obvious 
external  character  by  which  to  distinguish  them  as  a  group. 

Within  the  group,  the  species  from  which  its  name  is  taken  (because 
the  oldest  one  known)  is  most  easily  identified  in  the  adult  stage  from 
the  other  American  species  by  the  coarseness  of  the  sculpture  of  its 
bony  callosities  (pis.  9,  10),  and  in  the  early  juvenile  stage  by  its 
unique  coloration  (pi.  19).  The  greater  extension  of  the  plastral 
flap  anteriorly  beyond  the  carapace  is  also  a  noteworthy  feature. 


STEJXEGER:   AMERICAN    SOFT-SHELL   TURTLES  25 

Amyda  ferox1  (Schneider) 

Plates  5-10 

1783.— Tesludo  ferox  SCHNEIDER,  Naturg.  Schildkr.,  p.  330  (Savannah 
River,  Georgia;  type  in  Brit.  Mus.;  Dr.  A.  Garden,  collector)  (based 
on  Pennant,  Philos.  Trans.,  61,  pt.  1,  p.  266).— GMELIN,  Syst. 
Nat.,  1,  pt,  3,  1789,  p.  1039.— SCHOEPFF,  Naturg.  Schildkr.,  pt. 
5,  1795,  p.  102;  Hist.  Testud.,  pt.  5,  1795,  p.  88  (based  on  Pennant).- 
SHAW,  Gen.  Zool.,  3,  1802,  p.  64,  pi.  17.— LATREILLE,  Hist.  Nat. 
Rept,,  1,  1801,  p.  165  (based  on  Pennant).— DAUDIN,  Hist.  Nat. 
Rept.,  2,  1802,  p.  69  (based  on  Pennant). 
Trionyx  ferox  SCHWEIGGER,  Konigsberg.  Arch.  Naturw.  Math.,  1, 
1812,  pt.  3,  p.  285  (Carolina  and  Florida) ;  pt.  4,  p.  363;  Prodr.  Mon. 
Chelon.,  pt.  1,  1814,  p.  15.— MERREM,  Tent.  Syst.  Amph.,  1820, 
p.  20. — SAY,  Journ.  Acad.  Nat.  Sci.  Philadelphia,  ser.  1,  4,  pt.  2, 
1825,  p.  218  (part:  Carolina;  Georgia). — HARLAN,  Journ.  Acad. 
Nat.  Sci.  Philadelphia,  6,  Feb.  1827,  p.  32  (part:  many  of  the  rivers 
of  the  southern  states,  not  observed  to  exist  further  south  [north  ?] 
than  South  Carolina) ;  Medic.  Phys.  Res.,  1835,  p.  158.— Le  CONTE, 
Ann.  Lye.  Nat.  Hist.,  New  York,  3,  1830,  p.  93  (part:  Rivers  of 
Georgia  and  Florida,  north  to  Savannah). — GRAY,  Synops,  Rept., 
1831,  p.  45  (part);  Cat.  Tort.  Brit.  Mus.,  1844,  p.  49  (part);  Cat. 
Shield  Rept.  Brit.  Mus.,  pt.  1,  March  1856,  p.  68  (part);  HOL- 
BROOK,  North  Amer.  Herpet.,  ed.  1,  4,  1840,  p.  9,  pi.  1  (part: 
Savannah  and  rivers  emptying  into  northern  border  of  Gulf  of 
Mexico);  ed.  2,  2,  1842,  p.  11,  pi.  1  (part);  in  White's  Statistics  of 
Georgia,  1849,  Fauna  and  Flora,  p.  13  (Georgia).— STRAUCH, 
Mem.  Acad.  Sci.  St.  Petersbourg,  ser.  7,  5,  no.  7,  1862,  p.  173 
(part);  8,  no.  15,  1865  p.  122.  — BOULENGER,  Cat.  Chel.  Brit. 
Mus.,  1889,  p.  259  (Georgia,  Louisiana).— DITMARS,  Rept.  Book, 
1907,  p.  74,  pi.  26,  lower  fig.  (Georgia,  Florida,  Louisiana). — SIE- 
BENROCK,  Sitzungsbr.  Akad.  Wiss.  Wien,  Math.  Nat.  CI.,  91,  pt. 
1,  Oct.  1902,  p.  829;  Zool.  Jahrb.  Suppl.,  10,  pt.  3,  1909,  p.  603 
(Georgia,  Florida  west  to  Louisiana);  Verh.  Zool.  Bot.  Ges.  Wien, 
73,  Aug.  1923,  p.  181.  — BRIMLEY,  Proc.  Biol.  Soc.  Washington, 
23,  1910,  p.  18  (Georgia:  MimsviUe;  Florida:  Orlando;  Belleaire; 
Green  Cove  Springs;  St.  Petersburg).— DECKERT,  Copeia,  No. 
54,  Feb.  17,  1918,  p.  31  (Jacksonville,  Duval  Co.,  Fla.).— FLOWER, 
Proc.  Zool.  Soc.  London,  ser.  A,  1937,  pt.  1,  Apr.  15,  pp.  16,  37 
(age:  25  years  +  ). —  POPE,  Turtles  of  the  United  States  and 
Canada,  1939,  p.  303,  pi.  figs.  94-97  (Southeastern  Atlantic  and 
Gulf  Coastal  Plain). 

1  Ferocious,  the  specific  name  evidently  refers  to  the  following  sentence  in  Dr.  Garden's 
original  description:  "As  the  animal  is  very  fierce,  when  it  is  attacked  or  disturbed,  it  often 
raises  itself  on  its  legs,  and  will  leap  forward  to  bite  its  disturber  or  enemy,  which  it  does  with 
great  fury  and  violence". 


26  bulletin:  museum  of  comparative  zoology 

Amyda  ferox  OKEN,  Lehrb.  Zool.,  2,  1816,  p.  348.— STEJNEGER 
and  BARBOUR,  Check  List  North  Amer.  Amph.  Rept.,  ed.  1,  1917, 
p.  124  (South  Carolina  to  Florida  and  Louisiana);  ed.  2,  1923,  p.  140; 
ed.  3,  1933,  p.  153;  ed.  4,  1939,  p.  171.— SHUFELDT,  Aquat.  Life, 
6,  1920,  p.  27  (Georgia  to  Florida  and  Louisiana;  habits). — LODING, 
Alabama  Mus.  Nat.  Hist.,  Paper  No.  5,  Sept.  1922,  p.  47  (Fig 
Island,  Mobile  Co.,  Ala.).— PRATT,  Man.  Vert.  United  States, 
1923,  p.  250.— WRIGHT,  Ecology,  7,  Jan.  1926,  p.  84,  pi.  5,  fig.  3 
(Okefinokee  region,  Ga.).— CORRINGTON,  Copeia,  1927,  No.  165, 
Dec.  23,  p.  101  (Pascagoula  Swamp,  near  Biloxi,  Harrison  Co., 
Miss.).— PICKENS,  Copeia,  1927,  No.  165,  Dec.  23,  p.  43  (South 
Carolina).— JORDAN,  Man.  Vert.  Northeast  U.S.,  (13  ed.)  1929,  p. 
255  S.C.  to  Fla.  &  La.— CONANT,  Bull.  Antiven.  Inst.  America,  4, 
No.  3,  Dec.  1930,  p.  63  (Florida:  Seminole;  15  m.  E.  of  Sarasota;  18 
m.  S.  of  Ft.  Myers).— HALTOM,  Alabama  Mus.  Nat.  Hist.,  Pap. 
No.  11,  1931,  p.  141  (pi.  39)  (Alabama  part,  Mobile  Co.,  Fig  IsL). — 
VAN  HYNING,  Copeia,  1933,  No.  1,  Apr.  5,  p.  7  (Alachua  Co., 
Fla.,  moderately  common). — DeSOLA  and  ABRAMS,  Copeia, 
1933,  No.  1,  Apr.  3,  p.  12  (Okefinokee  Swamp,  Ga.).— ALLEN, 
Amer.  Mus.  Novit.,  No.  542,  June  20,  1932,  p.  20  (Mississippi:  Han- 
cock Co.).— RUST,  Blatt.  Aquar.  Terrarienk.,  45,  1934,  sep.  p.  12. 

Platypeltis  ferox  FITZINGER,  Syst.  Rept.,  1843,  p.  30.— AGASSIZ, 
Contr.  Nat.  Hist.  United  States,  1,  1857,  p.  401  (southern  states, 
Georgia  to  western  Louisiana). — GRAY,  Proc.  Zool.  Soc.  London, 
1869  (p.  214);  1873  (p.  58)  (part).— TRUE,  Fish.  Industr.  United 
States,  sect.  1,  1884,  p.  152.— BAUR,  Proc.  Amer.  Philos.  Soc,  31, 
1893,  p.  220— LONNBERG,  Proc.  U.  S.  Nat.  Mus.,  17,  1894,  p. 
317  (southern  Florida).— WRIGHT  and  BISHOP,  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  vol  67,  1915,  p.  119,  pi.  1,  figs.  1,  2,  4;  pi.  2,  fig.  6 
(Okefinokee  Swamp,  Georgia). 

Aspidonectes  ferox  COPE,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  1875,  p.  51 
(Georgia  to  western  Louisiana), — TRUE,  in  H.  S.  Thompson,  South 
Carolina,  1883,  p.  238  (South  Carolina);  Bull.  U.  S.  Nat.  Mus.,  No. 
24,  1883,  p.  29  (part:  Palatka,  Putnam  Co.,  Fla.;  Charleston,  S.  C). 
—DAVIS  and  RICE,  Illinois  State  Lab.  Nat.  Hist.  Bull.  No.  5, 
1883,  p.  52  (Georgia  to  western  Louisiana).— SHUFELT,  Rep.  U.S. 
Nat.  Mus.  1892  (1893)  p.  32  (cast  U.S.N.M.  No.  8899).— BAUR, 
Amer.  Natural.,  22,  1888  (p.  1121).— COKER,  Bull.  North  Carolina 
Geol.  Surv.,  14,  1906,  p.  66  (North  Carolina,  introduced  ?).— GOFF 
and  GOFF,  Copeia,  1935,  No.  3,  Oct.  15,  p.  156  (Griffin,  Lake  Co., 
Fla.;  incubation). 

1788.— Testudo  mollis  LACEPEDE,  Hist.  Nat.  Quadr.  Ovip.  Serp.,  1,  Synops. 
Meth.  Quadr.  Ovip.,  tab.  between  pp.  618  and  619  (based  on  Pen- 
nant).— BONNATERRE,  Tabl.  Enc.  Meth.  Erpet.,  1789,  p.  25. 


stejneger:  American  soft-shell  turtles  27 

1795.— Testudo  (ferox  ?)  verrucosa  SCHOEPFF,  Naturg.  Schildkr.,  pt.  5,  p.  105 
(based  on  Bartram's  Trav.  North  and  South  Carolina,  1791,  p.  177) 
(type  locality,  eastern  Florida);  Hist.  Testud.,  pt.  5,  1795,  p.  90. 

1801.— Testudo  bartrami  DAUDIN,  Hist.  Nat.  Rept.,  2,  p.  74  (based  on  Bar- 
tram,  Voy.  Amer.  Septentr.,  vol.  1,  p.  307).— HARPER,  Amer.  Mid- 
land Natural.,  23,  No.  3,  May  1940,  p.  717  (Halfway  Pond,  Putnam 
Co.,  Florida,  "restricted  type  locality"). 
Trionyx  bartrami  GEOFFROY-ST.  HILAIRE,  Ann.  Mus.  Hist.  Nat. 
Paris,  14,  Aug.  1809  (p.  18).— Le  CONTE,  Ann.  Lye.  Nat.  Hist. 
New  York,  3,  1830,  p.  96  (St.  John's  River,  Fla.). 

1809  1— Trionyx  carinatus  GEOFFROY-ST.  HILAIRE,  Nouv.  Bull.  Soc. 
Philom.  Paris,  1,  No.  22,  July  1809,  p.  365  (type  locality  unknown; 
type  in  Paris  Mus.);  Ann.  Mus.  Hist.  Nat.  Paris,  14,  Aug.  1809,  p. 
14,  pi.  4. 

1809.— Trionyx  georgianus  GEOFFROY-ST.  HILAIRE,  Nouv.  Bull.  Soc. 
Philomat.  Paris,  1,  No.  22,  July  1809,  p.  367  (substitute  name  for 
T.  ferox  "Pennant"). 

1809.— Trionyx  georgicus  GEOFFROY-ST.  HILAIRE,  Ann.  Mus.  Hist.  Nat- 
Paris,  14,  Aug.  1809,  p.  17  (variant). 

1812  ?.— Trionyx  brongniarti  SCHWEIGGER,  Konigsberg.  Arch.  Naturw- 
Math.,  1,  p.  288  (substitute  name  of  Trionyx  carinatus);  Prodr.  Mom 
Chelon.,  1814,  p.  18. 

1835. — Gymnopus  spiniferus  DUMERIL  and  BIBRON,  Erpet.  Gen.,  2,  p.  477 
(part:  Rivers  of  Georgia  and  Florida). 

1835.— Trionyx  harlani  "Bell"  HARLAN,  Medic.  Phys.  Research,  p.  159 
(type  locality,  East  Florida;  type  in  "Mus.  of  Bell;  Lond.").— 
DeKAY,  Zool.  New  York,  Rept.,  1842,  p.  7  (East  Florida). 

Types.  The  type  of  Testudo  ferox  is  still  extant  in  the  British 
Museum.1  The  species  was  discovered,  described  and  illustrated  by  an 
American;  his  descriptions  and  illustrations  were  published  by  one  of 
the  most  outstanding  English  zoologists  of  the  period  in  the  foremost 
scientific  journal  of  Europe;  Linnaeus  himself  was  simultaneously 
notified  of  its  discovery.  Yet,  for  twelve  years  it  remained  without  a 
systematic  name,  until  a  German  bestowed  on  it  the  name  by  which  it 
is  specifically  known.  This  unusual  nomenclatorial  history  and  the 
inaccessibility  of  the  original  description  to  most  zoologists  justify  its 
reproduction  here  with  a  brief  account  of  the  related  circumstances. 

Dr.  Alexander  Garden,  of  Charleston,  who  was  an  ardent  student  of 

1  According  to  Shaw  (Gen.  Zool.,  3,  pt.  1,  1802),  the  specimen  was  already  then  in  the  British 
Museum.  In  1831  Gray  (Syn.  Rept.,  p.  46)  confirms  that  statement  and  adds  that  Pennant 
gave  the  type  specimen  to  the  Royal  Society.  In  1844  and  1856  he  records  it  as  the  "Specimen 
described  and  figured  by  Pennant,  (restuffed)." 


28  bulletin:  museum  of  comparative  zoology 

the  natural  history  of  the  Carolinas,  became  possessed  of  an  adult 
specimen  of  the  first  softshell  turtle  reported  from  America.  On 
December  24,  1770,  he  wrote  to  his  friend  Mr.  John  Ellis  in  London 
as  follows:1 

"I  have  one  or  two  things  which  I  think  will  please  him  [Thomas  Pen- 
nant] ;  he  shall  have  them  by  one  of  our  spring  ships ;  one  of  them  is  a 
species  of  Turtle,  as  yet  nondescript.  It  is  amazing  how  Catesby 
omitted  this.  It  is  found  in  abundance  in  the  Savannah  river,  in 
Altamaha,  and  East  Florida.  It  is  a  fresh  water  animal,  grows  to  a 
great  size,  and  is  as  delicate  as  the  Green  Turtle,  having  a  large 
leathery  cover  over  its  back,  and  a  head  very  like  a  Mole.  I  intend  to 
send  a  copy  of  my  account  of  this  animal  to  Mr.  Pennant  for  his 
American  Zoology,  and  if  I  can  get  a  drawing  of  it  copied,  I  will  send 
him  that.  If  I  can  obtain  another  Turtle,  I  shall  send  you  one  stuffed. 
It  has  a  relation  to  the  first  species  of  Linnaeus's  last  edition  of  the 
Systema  Naturae."2 

Early  next  spring  he  kept  his  promise  and  sent  Pennant  the  descrip- 
tion as  well  as  a  preserved  specimen.  Another,  better  specimen,  he 
forwarded  at  the  same  time  to  Mr.  Ellis  with  a  drawing  made  from 
life.  This  specimen  with  the  drawing  and  the  letter  Pennant  laid  be- 
fore the  Royal  Society  in  London  on  May  2,  1771,  in  the  Philosophical 
Transactions  of  which,  vol.  61,  they  were  published.  Dr.  Garden  does 
not  state  exactly  whence  the  specimens  came.  In  his  letter  he  says  that 
"they  are  not  commonly  got  here  in  Charles-town,  though  by  chance 
this  last  summer,  I  had  two  sent  me",  but  as  he  mentions  Savannah 
and  Altamaha  rivers,  it  may  be  inferred  that  the  two  turtles  were  sent 
him  from  one  of  those  rivers,  or  one  from  each  of  them.  One  of  the 
specimens,  "the  specimen  described  and  figured  by  Pennant  (re- 
stuffed)",  according  to  Gray  is  the  type  in  British  Museum. 

Dr.  Garden's  original  description  follows : 

[p.  268]  "They  are  found  in  large  quantities  in  Savannah  and  Alta- 
maha rivers;  and  I  have  been  told  that  they  are  very  common  in  the 
rivers  in  East  Florida. 

They  grow  to  very  large  sizes,  though  the  largest  that  ever  I  heard 
of  was  seventy  pounds. 

The  Turtle,  which  I  now  have  by  me,  weighs  twenty  pounds;  and 
probably,  when  I  first  got  it,  it  might  have  weighed  from  twenty-five 
to  thirty  pounds,  as  I  have  observed  that  it  has  grown  poorer  every 

1  Smith,  J.  E. :  A  Selection  of  the  Correspondence  of  Linnaeus  and  other  Naturalists.  London, 
1821,  1,  p.  580. 

2  Linnaeus,  Systema  Naturae,  ed.  12,  1,  1766,  p.  350  (Testudo  coriar.en\. 


stejneger:  American  soft-shell  turtles  29 

day.  I  have  had  it  now  near  three  months,  and  I  never  could  observe 
that  it  has  eaten  any  thing  that  has  been  given  it,  though  a  variety  of 
things  have  been  tried. 

It  is  twenty  inches  long  from  one  end  of  the  shell  or  covering  to  the 
other,  and  fourteen  inches  and  a  half  bread.  The  colour  of  this  shell 
or  covering,  in  general,  is  dark  brown,  with  a  greenish  cast. 

The  middle  part  is  hard,  strong,  and  bony;  but  all  round  the  sides, 
especially  towards  the  tail  and  hindermost  part,  it  is  cartilaginous, 
soft  and  pliable,  resembling  thick  tanned  sole-leather,  yielding  very 
easily  to  any  force  in  any  direction  whatever,  but  thick  enough  and 
strong  enough  to  defend  the  animal  from  any  injury.  All  the  hind 
part  of  the  back  is  full  of  oblong  smooth  knobs;  and  the  fore  part, 
just  where  it  covers  the  head  and  neck,  is  studded  full  of  large  knobs. 
The  under  side  of  this  plate  is  very  [p.  269]  beautiful,  of  a  lively  whitish 
colour,  interspersed  with  innumerable  very  fine  ramifications  of  blood 
vessels,  running  from  the  margin  of  the  plate  into  larger  and  larger 
branches,  until  the  sight  of  them  is  at  once  lost  by  their  entering  the 
body  of  the  animal. 

The  under,  or  belly  plate,  or  rather  sternum,  is  of  a  fair  whitish 
colour,  and  extended  forward  two  or  three  inches  more  than  the  back 
plate,  so  that  the  head  rests  on  it  very  conveniently.  The  hind  part 
of  this  plate  is  hard  and  bony,  shaped  very  much  like  a  man's  riding 
saddle,  with  two  pieces  for  the  thighs  to  rest  on.  The  fore  part  of  the 
plate  is  pliable  and  cartilaginous. 

The  head  is  somewhat  triangular  and  attenuated,  rather  apparently 
small  for  the  animal,  but  growing  gradually  larger  towards  the  neck, 
which  is  thick  and  long,  and  easily  extended  out  (neck  of  the  present 
subject  was  thirteen  inches  and  a  half  long)  to  a  great  length,  or  drawn 
back  again  under  the  shell  or  plate. 

The  eyes  are  placed  in  the  fore  and  upper  part  of  the  head,  near  to 
one  another,  having  pretty  large  loose  palpebrae.  The  pupil  is  small 
and  lively,  surrounded  by  a  lemon-coloured  iris,  perfectly  round,  and 
giving  much  life  and  fire  to  the  eyes.  When  danger  approaches,  or 
when  it  goes  to  sleep,  it  covers  its  eyes,  by  bringing  the  inner  and 
loose  part  of  the  lower  palpebrae  over  its  eye,  like  a  membrana  nictitans. 

The  upper  lip  and  under  lip  are  both  large,  but  especially  the  upper. 
The  mandibula  are  both  entire,  each  being  one  entire  bone  all  round, 
of  the  same  shape  as  the  mouth. 

[p.  270]  The  nostrils  are  the  most  singular  part,  being  a  cartilagi- 
nous production  of  at  least  three  quarters  of  an  inch,  beyond  the  upper 
and  fore  angle  or  point  of  the  upper  lip,  perforated  with  two  apertures 


30  bulletin:  museum  of  comparative  zoology 

reaching  back  and  opening  into  the  roof  of  its  mouth,  having  a  smooth 
septum  but  fimbriated  upon  each  side.  This,  at  first  sight,  in  some 
manner  resembles  the  snout  of  the  mole;  but  it  is  tender,  thin  and 
transparent,  and  cannot  be  intended  for  digging  in  the  earth  or  land. 

The  arms  are  thick  and  strong,  consisting  of  three  distinct  joints, 
viz.  the  upper,  the  fore  arm,  and  hand.  The  hands  have  each  five 
fingers,  of  which  the  three  first  are  shorter  and  stronger,  and  furnished 
with  strong  nails,  or  rather  claws.  The  two  last  fingers  have  more 
joints,  but  are  smaller,  and,  instead  of  being  furnished  with  claws,  are 
covered  with  the  membrane,  which  is  extended  even  beyond  their  ex- 
tremities. Towards  the  back  or  hind  part,  there  are  two  spurious 
fingers,  which  just  serve  to  support  the  membrane  when  extended. 
The  upper  side  of  these  arms  and  hands  are  covered  with  a  wrinkled 
loose  skin,  of  a  dusky  greenish  colour.  The  legs  consist  of  the  same 
number  of  joints,  and  have  the  same  number  of  toes  as  there  are  fingers 
on  the  fore-feet,  and  these  are  furnished  with  nails  in  the  same  manner, 
only  there  is  but  one  spurious  toe.  Both  the  fore  and  hind  legs  are 
thick,  strong,  and  muscular;  and  as  the  animal  is  very  fierce,  when  it  is 
attacked  or  disturbed,  it  often  raises  itself  on  its  legs,  and  will  leap 
forward  to  bite  its  disturber  or  enemy,  which  it  does  with  great  fury 
and  violence. 

[p.  271]  They  are  likewise  very  strong,  and  of  a  lively  whitish 
colour,  because  they  are  generally,  if  not  always,  covered  with  the 
upper  plate,  which,  as  I  said  before,  is  extended  a  great  way  behind. 

The  tail  is  large  and  thick,  and  generally  as  long  as  the  hind  part  of 
the  upper  plate.  The  anus  is  placed  about  an  inch  from  the  extremity 
of  the  tail  on  the  inside. 

The  turtle,  from  which  these  characters  were  taken,  was  a  female; 
after  she  came  into  my  possession  she  laid  fifteen  eggs,  and  about  the 
same  number  were  taken  out  of  the  belly  when  she  died.  The  eggs  were 
nearly  an  inch  diameter,  and  perfectly  spherical. 

It  is  esteemed  very  good  eating,  and  said  by  many  to  be  more 
delicate  than  the  green  turtle." 

On  June  20,  1771,  Dr.  Garden  wrote  a  letter  in  Latin  to  Linnaeus 
which  is  translated  as  follows  :l 

"I  have  described  and  have  lately  sent  to  our  friends  Ellis  and 

Pennant,  a  new  and  very  rare  species  of  river  Tcstudo,  known  here 

by  the  name  of  the  Softshelled  Turtle,  because  the  covering  of  its 

back,  especially  towards  the  sides,  is  of  a  softish,  leathery,  very 

1  Smith,  op.  cit.  p.  336. 


stejneger:  American  soft-shell  turtles  31 

flexible  substance.    This  animal  is  found  in  the  larger  fresh-water 
rivers  of  East  Florida,  Georgia  and  South  Carolina." 
Dr.  Garden  probably  was  hoping  that  Linnaeus  might  have  hastened 
to  supply  the  still  missing  nomcn  triviale  as  he  did  five  years  earlier 
with  Garden's  no  less  startling  discovery  of  Siren  lacertina. 

It  will  be  noted  that  Dr.  Garden  himself,  in  the  letter  to  Linnaeus, 
identified  the  new  discovery  with  the  softshell  occurring  in  East 
Florida,  and,  subsequently,  naturalists  applied  the  name  ferox  indis- 
criminately to  all  specimens  from  North  America.  Therefore,  when 
Lesueur  in  1827  described  T.  spinifcrus  as  a  distinct  species  and  his 
great  countrymen  Cuvier  and  Dumeril  declared  it  to  be  only  the  young 
of  T.ferox,  the  identity  of  the  two  names  was  generally  accepted,  even 
by  Holbrook  (1842)  and  Gray  (1856). *  Not  until  1857  when  Agassiz 
demonstrated  the  distinctness  of  spinifcrus  did  it  become  generally 
recognized,  although  even  Boulenger's  treatment  (1889)  shows  that 
the  true  characters  of  T.  ferox,  as  represented  by  the  type  specimen, 
were  not  completely  understood. 

The  type  specimen  in  British  Museum  was  examined  by  Dr.  Georg 
Baur  on  September  7,  1888,  as  closely  as  the  circumstances  then  per- 
mitted. He  noted  particularly  its  "sehr  rauhe  Ornament"  (very  rough 
sculpture),  and  "Keine  Spines,  sondern  Tuberkel"  (no  spines,  but 
tubercles)  evidently  as  compared  with  spinifcrus;  "Schadel  vom  Typus 
des  Exempl.  von  Lucas.  Vorderer  Theil  stark  beschadigt,  stark  vorn 
abfallend.  Unterkief.  ganz  von  jenem  Typus.  Jugale  mit  minim,  unt. 
Fortsaz."  (Skull  of  the  type  of  Lucas'  specimen.  Front  part  much 
injured,  greatly  inclining  anteriorly.  Lower  jaw  entirely  of  that  type. 
Jugal  with  minimal  lower  process).  He  sums  up  thus:  "Resultat :  Type 
von  Platy pcltis  ferox     Exemplar  von  Lucas." 

Lucas'  "exemplar"  is  U.S.N.M.  8899  (tintag  read  upside  down  6688 
by  Baur  in  his  notes)  which  Baur  had  been  studying  with  F.  A.  Lucas, 
then  curator  of  the  division  of  comparative  anatomy  in  the  National 
Museum.  No.  8899  is  a  fine  disarticulated  skeleton  still  in  the  Museum, 
of  approximately  the  same  size  as  Dr.  Garden's  type,  and  was  col- 
lected by  Professor  S.  F.  Baird,  April  1877,  in  the  St.  John's  River, 
Florida,  probably  not  far  from  Jacksonville.  A  fine  plaster  cast 
painted  by  Schindler  from  his  color  sketch  of  the  living  specimen  is 
on  exhibition  in  the  Museum.  A  photograph  of  it  was  published  in 
the  report  of  the  Museum  for  1882,  pi.  32. 

1  For  some  unexplained  reason,  however,  Dumeril  and  Bibron  ignored  the  priority  rights  of 
ferox  and  called  the  combination  which  included  the  South  Carolina  to  Florida  records  Gymnopus 
iferus.   Possibly  as  a  disguised  protest  against  Geoffroy's  inefficient  substitution  of  Trionyx 
for  Schweigger's  earlier  Emyda. 


32 


bulletin:  museum  of  comparative  zoology 


Thanks  to  the  authorities  of  the  British  Museum  I  have  been  per- 
mitted to  examine  the  type  and  can  confirm  completely  Dr.  Baur's 
result  as  demonstrated  by  the  photographs  side  by  side  (pi.  5)  of  the 
two  specimens  and  the  measurements  in  table  1. 

Type  of  Trionyx  harlani.     In  the  Medical  and  Physical  Researches 
published  by  him  in  1835  Dr.  Richard  Harlan  includes  on  p.  159  the 
description  of  a  Trionyx  as  follows : 
"Trionyx  Harlani 
Trionyx  Harlani,  Bell,  Monogr.  Test,  pi. 
Char.     Body  more  ventricose,  soft  portions  of  the  shell  less  exten- 
sive than  in  the  other  species.   In  general  appearance  ap- 
proaching more  to  the  genus  Emys. 

Inhabits  East  Florida.  Mus.  of  Bell,  Lond." 
I  have  not  been  able  to  locate  any  such  plate  in  Bell's  Monograph 
of  the  Testudinata.  Bell,  in  correspondence,  may  have  indicated  that 
he  had  in  his  collection  a  specimen  from  East  Florida  which  he  intended 
to  figure  in  his  unfinished  Monograph  under  the  above  name,  but  there 
is  no  such  plate  among  the  unedited  ones  published  in  1872  under  the 
title:  "Tortoises,  Terrapins  and  Turtles",  neither  have  I  found  any 
record  of  what  became  of  Bell's  specimen.  J.  E.  Gray  in  the  introduc- 
tion of  his  catalogue  of  the  Tortoises  in  British  Museum,  notices  that 
"the  specimens  presented  by  ....  Thomas  Bell,  Esq.  [may  be  re- 
garded] as  the  types  of  the  species  described  in  his  various  papers,  and 

in  his  very  beautiful  Monograph  of  the  Testudinata [by]  Dr. 

Richard  Harlan,  and  Messrs.  Edward  and  Henry  Doubleday,  as  the 
types  of  the  North-American  species  described  by  Say,  Harlan,  and 
others,"  but  he  has  no  reference  to  any  softshell  from  Florida,  nor  is 
there  in  any  of  the  later  catalogs. 

External  characters.  While  the  skull  characters  are  only  available 
in  dubious  and  critical  cases,  normal  specimens  of  A.  fcrox  within  the 
group  are  not  difficult  to  identify,  the  older  ones  by  the  coarseness  of 
the  sculpture  of  the  bony  carapace;  the  very  young  ones  by  their 
unique  coloration. 

The  coarseness  of  the  sculpture  of  the  bony  carapace  of  A.  ferox  is 
not  confined  to  the  general  network  of  vermiculations  of  the  surface, 
but  it  is  commonly  specialized  into  a  series  of  more  or  less  prominent 
longitudinal  welts.  The  difference  in  relative  size  and  pattern  of  the 
pits  and  ridges  which  constitute  the  character  of  the  "sculpture"  is 
difficult  to  describe,  but  a  comparison  of  the  samples  figured  (pis.  7, 
9,  10)  explains  it  better  than  words. 

The  unique  coloration  of  the  hatchlings  and  the  very  young  speci- 


stejneger:  American  soft-shell  turtles 


33 


mens  constitutes  the  most  obvious  and  characteristic  feature  of  the 
species,  but  unfortunately  disappears  with  age.  In  all  the  other  species 
the  young  are  of  a  more  or  less  pale  olive  or  tawny  ground  color,  either 
uniform  or  marked  with  dusky  or  blackish  specks  of  varying  shape, 
but  mostly  round  dots,  which  when  larger  assume  the  form  of  ocelli 
with  a  lighter  center,  combined  with  a  nearly  uniform  white  plastron 
(except  for  the  pinkish  tinge  due  to  the  fine  blood  vessels  shining 
through  in  the  living  specimens).  The  prevailing  feature  is  the  dark- 
ness and  the  saturation  of  the  pigmentation,  the  big  dark  blotches  on 
the  carapace,  and  the  dark  slate  gray  underside,  in  combination  with 
the  strong  contrast  of  the  light  pattern  on  the  head  and  the  margin  of 
the  carapace,  detailed  description  of  which  will  be  given  below. 

The  anterior  flap  of  the  leathery  plastron  is  longer  and  less  circular 
than  in  the  other  species,  often  extending  a  considerable  distance  for- 
ward beyond  that  of  carapace,  but  it  appears  to  vary  individually  and 
is  difficult  of  precise  definition  because  of  absence  of  suitably  fixed 
points  from  which  to  measure. 

The  shape  of  the  outline  of  the  disk,  however,  is  somewhat  different 
in  the  }roung  of  A.  ferox,  in  as  much  as  it  is  less  circular  than  in  the 
other  species,  as  will  appear  from  table  3  which  presents  the  measure- 
ments of  five  hatohlings  of  the  same  brood.  Similar  proportions  are 
shown  by  one  collected  by  Dr.  Francis  Harper  in  the  Okefinokee 
Swamp  (U.S.N.M.  84603),  viz.  length  of  leathery  carapace  39.5  mm. 
and  width  32.5  mm.  It  is  also  well  illustrated  in  the  photograph  of 
No.  61087,  pi.  17,  fig.  a. 

Table  3 

Amyda  ferox,  pullus 

Florida:  Polk  Co.,  Auburndale 


mm. 


Length  of  leathery  disk 
Width  of  leathery  disk     mm. 
Height  of  body 


mm. 


61083 


38.0 
33.5 

12.5 


61084 


40.5 
35.5 
12.0 


61085 


40.0 
34.0 
12.5 


61086 


41.0 
34.0 
12.0 


61087 


45.0 
37.0 
14.0 


Average 


40.9 
34.8 
12.6 


Size.     Amyda  ferox  is  apparently  the  largest  of  the  North  American 
softshell  turtles.    Authentic  measurements  of  large  carapaces  are  few 


34  bulletin:  museum  of  comparative  zoology 

and  those  available  are  of  dubious  value,  due  to  uncertainty  as  to 
identification,  condition  of  specimen  when  measured,  and  method  of 
measurement  whether  in  a  straight  line  or  along  the  curvature  of  the 
shell.  The  largest  Agassiz  "had  ever  seen  or  heard  of"  was  one  from 
Natchez  "which  measured  eighteen  inches  and  a  half  [470  mm.]  from 
the  front  to  the  hind  margin  of  the  carapace"  (Contrib.  Nat.  Hist. 
U.  S.,  vol.  1,  p.  401).  The  largest  specimen  now  in  the  National 
Museum  is  an  old  skin,  with  the  skull  in  (U.S.N.M.  No.  38123  from 
"Florida"  which  measures  about  17  inches  along  the  curvature  (430 
mm.).  Its  zygomatic  width  is  68  mm.  The  plaster  cast  of  No.  8899 — 
(in  exhibition  series)  is  438  mm.  long;  over  the  curvature  it  measures 
460  mm.  (18  inches);  zygomatic  width  67  mm. 

But  larger  specimens  may  exist,  or  have  been  living  in  Florida  not 
long  ago.  The  National  Museum  has  a  series  of  16  weathered  skulls 
picked  up  by  Dr.  E.  A.  Mearns  near  Kissimee  about  the  beginning  of 
this  century,  14  of  them  larger  than  that  of  the  18  inch  specimen  men- 
tioned above  (No.  8899).  The  basicranial  length  of  this  one  is  92  mm., 
the  corresponding  dimension  of  the  14  Kissimee  skulls  range  from  95 
to  114  mm.  (aver.  103)  with  the  zygomatic  width  varjing  between 
65  and  80  mm.  (aver.  73). 

Abnormal  alveolar  surfaces 

This  series  of  16  skulls  of  evidently  very  old  specimens  shows  an 
extraordinary  development  which  deserves  special  attention. 

In  the  normal  skulls  of  the  species  in  all  our  specimens  with  a  basi- 
cranial length  below  90  mm.  the  lateral  outline  of  the  snout  anterior 
to  the  orbit  tapers  towards  the  end  in  a  fairly  straight  line,  and  the 
narrow  maxillary  alveolar  surface  follows  almost  parallel,  as  in  the 
other  species  of  the  ferox  group,  irrespective  of  sex  and  age. 

However,  in  the  series  of  16  Mearns  skulls  with  basicranial  length 
above  84  mm.  we  find  two  different  styles  of  snout  outline  and  alveolar 
surface  as  recorded  in  table  4. 

For  the  illustration  of  the  extremes  of  the  width  of  the  alveolar  sur- 
face and  the  outline  of  the  maxilla  it  is  only  necessary  to  refer  to  plate 
6.  While  greatly  reduced  the  figures  convey  the  difference  between  the 
two  series  as  the  figures  are  of  the  same  relative  size. 

Evidently  the  abnormal  development  of  the  maxilla  as  represented 
by  plate  6,  figs.  3  and  4  are  due  to  old  age  since  we  do  not  find  it  in  the 
smaller  and  younger  specimens.  It  at  once  recalls  similar  conditions 
in  some  Chinese  species  of  the  genus,  upon  which  Father  Heude,  in 


stejneger:  American  soft-shell  turtles 


35 


Table  4 
Alveolar  width  and  outline  of  snout  in  a  series  of  very  old  A.  ferox. 


Snout  outline  straight 

Snout  outline  strongly 

convex 

Greatest 

Greatest 

U.S.N.M. 

alveolar 

Basicranial 

Basicranial 

alveolar 

U.S.N.M. 

No. 

width 

length 

length 

width 

No. 

029459 

7.5  mm. 

114  mm. 

108  mm. 

21.5  mm. 

029464 

107  mm. 

15.5  mm. 

029458 

106  mm. 

14.0  mm. 

029454 

105  mm. 

14.0  mm. 

029470 

105  mm. 

16.0  mm. 

029457 

104  mm. 

18.0  mm. 

029463 

104  mm. 

11.5  mm. 

029451 

101  mm. 

15.0  mm. 

029450 

101  mm. 

11.5  mm. 

029460 

100  mm. 

14.0  mm. 

029455 

029456 

7.5  mm. 

100  mm. 

97  mm. 

11.5  mm. 

029453 

95  mm. 

9.5  mm. 

029452 

029475 

6.8  mm. 

92  mm. 

84  mm. 

12.0  mm. 

029462 

1880, 1  based  the  description  of  numerous  new  genera  and  species.  I 
need  only  refer  to  his  pictures  of  Caelognathus  novemcostatus,  pi.  5; 
Tortisternum  novemcostatum,  pi.  G;  Cinctisternum  bicinctum,  pi.  9;  and 
especially  Ceramopelta  latirostris,  pi.  7.  Notwithstanding  the  fact  that 
he  did  not  obtain  any  young  specimens  with  these  characteristics 
Heude  regarded  them  as  specific  or  generic  differences.  Boulenger  in 
cataloguing  the  softshell  turtles  in  the  British  Museum  (1889,  p.  243) 
was  "unable  to  find a  single  young  specimen  with  the  molar- 
like alveolar  surfaces"  but  having  "found  in  three  species,  viz.  T. 
triunguis  (Africa),  T.  cartilagineus  (E.  Indies),  and  T.  sinensis  (China) 


i  Mem.  Hist.  Nat.  Emp.  Chinois,  1,  pp.  1-38,  pis.  1-9. 


36  bulletin:  museum  of  comparative  zoology 

examples  of  the  two  types,  i.e.  on  the  one  hand  sharp-edged,  com- 
paratively narrow  jaws,  and  on  the  other  hand  broad  crushing  alveolar 
surfaces  nearly  meeting  on  the  median  line  in  front  of  the  choanae, 

in  specimens  which,  in  other  respects,  are  undistinguishable, 

[he]  arrived  at  the  conclusion  that  we  may  be  in  presence  of  a  case  of 
dimorphism  caused  by  a  difference  of  diet." 

This  idea  of  a  "dimorphism"  due  to  a  difference  of  diet  does  not  seem 
convincing.  That  it  is  not  the  result  of  old  age  alone  seems  obvious 
from  the  fact  that  so  many  old  skulls  of  the  same  size  and  presumably 
age  do  not  show  the  abnormality.  That  it  is  not  due  to  a  change  of 
diet  affecting  the  whole  population  seems  also  obvious  since  the  Kissi- 
mee  series  are  all  practically  from  the  same  locality.  That  the  differ- 
ence is  not  a  local  one  is  proved  by  a  weathered  skull  almost  identical 
with  Kissimee  No.  029462  of  the  above  table,  picked  up  by  Dr.  Francis 
Harper  in  the  Okefinokee  Swamp  (U.S.N.M.  No.  59727)  which  with 
a  basicranial  length  of  88  mm.  has  a  maxillary  alveolar  width  of  11 
mm.  The  fact  that  in  the  series  the  largest  skull  is  normal  and  the 
smallest  abnormal  coupled  with  the  other  fact  that  all  the  largest 
authentically  sexed  specimens  in  the  collection  are  female  and  normal, 
while  among  the  preserved  specimens  no  abnormal  male  has  been 
recorded,  suggest  that  the  difference  may  be  due  to  sex.  If  so,  does  that 
indicate  an  individual  preference  among  the  old  males  for  a  certain 
kind  of  diet? 

The  normal  skull.  As  repeatedly  noted  the  skull  of  A.  ferox  is  built 
on  the  same  plan  as  that  of  spinifera  and  cmoryi  and  differs  but  slightly 
in  the  proportionate  size  and  relation  of  the  various  bones,  but  the 
individual  variation  is  so  great  that  in  some  cases  it  is  even  difficult 
to  decide  to  which  species  an  isolated  skull  without  locality  record  be- 
longs. A.  ferox,  however,  may  generally  be  diagnosed  as  having  the 
narrowest  interocular  width,  the  shortest  distance  between  choanae 
and  intermaxillary  foramen,  and,  in  the  medium-sized  skulls,  the 
narrowest  maxillary  alveolar  surface.  It  also  averages  the  greatest 
length  from  orbit  to  tympanic  cavity  combined  with  the  shortest  and 
weakest  mandibular  symphysis.  The  interorbital  space  is  usually  less 
in  width  than  one  third  of  the  longest  diameter  of  the  temporal  fossa, 
while  in  the  others  it  is  usually  more  than  one  third.  The  anterior 
processes  of  the  prefrontals  as  a  rule  are  longer  and  slenderer  and  the 
angle  projecting  into  the  posterior  border  of  the  nasal  fossa  is  conse- 
quently more  acute.  In  the  older  and  younger  specimens  these  differ- 
ences become  more  obscure  or  may  be  entirely  obliterated. 

In  a  series  of  skulls  of  this  group  a  rather  common  feature  may  be 


stejneger:  American  soft-shell  turtles  37 

noted  in  the  larger  specimens  of  A.  ferox.  While  in  the  palate  of  the 
other  species  the  maxillaries  are  in  contact  practically  the  whole  length 
between  the  choanae  and  the  intermaxillary  foramen,  they  may  be  en- 
tirely or  partly  separated  by  the  vomer  which  in  many  skulls  may  be 
seen  as  a  fork  enclosing  the  posterior  end  of  the  foramen  as  shown  in 
all  the  figures  on  pi.  6  (very  large  specimens). 

Coloration.  Old  specimens  of  this,  as  well  as  the  other  species,  show 
but  little  of  the  characteristic  normal  coloration  and  pattern  of  the 
species.  In  A.  ferox  more  or  less  faint  remnants  of  the  large  brownish 
blotches  on  the  carapace  may  be  made  out  on  the  generally  dingy 
"Isabella"  colored  ground,  or  disappear  in  the  blackish  or  brownish 
variations  of  the  latter.  But  as  previously  indicated,  the  very  young 
ones  display  a  distinctive  and  peculiar  color  and  pattern,  unknown 
until  figured  by  Ditmars  (1907)  and  described  by  Wright  and  Funk- 
houser  (1915).  Some  of  the  specimens  received  by  the  National 
Museum  were  either  alive  or  freshly  killed  and  their  descriptions  with 
reference  to  a  standard  color  nomenclature  (Ridgway's  Nomenclature 
of  Colors  for  Naturalists,  1886)  were  made: 

U.S.N.M.  No.  61087.  Auburndale,  Polk  Co.,  Florida,  collected  by 
N.  R.  Wood,  summer  of  1918.  Pullus.  Length  of  leathery  carapace, 
45  mm.  Leathery  plastron  extending  anteriorly  beyond  carapace  6  mm. 
Iris  pale  silver  gray  with  a  horizontal  black  bar. — Upper  surface  of 
carapace  tawny  olive  (Ridgway,  pi.  iii,  fig.  17)  with  dusky  (dark 
sepia)  spots  and  a  narrow  well-defined  outer  edge  of  bright  ochraceous 
(R.  v,  7)  in  strong  contrast;  underside  of  carapace  and  plastron  slate- 
gray  (R.  ii,  5)  with  scattered  clay-colored  spots  on  the  former  and  the 
anterior  edge  of  plastron,  the  outer  edge  of  disk  narrowly  ochraceous, 
as  above,  though  less  bright;  upper  side  of  head,  neck  and  legs  olive 
(R.  iii,  9)  with  clay-colored  marblings;  from  anterior  angle  of  eyes  to 
middle  of  proboscis  an  inverted  Y-shaped,  pale  clay-colored  figure,  the 
fork  situated  halfway  between  eye  and  base  of  proboscis;  side  of  head 
olive  with  a  wide  well-defined  angular  band  of  yellowish  buff  extending 
from  posterior  corner  of  eye  to  base  of  lower  jaw;  another  similar 
band,  but  slightly  broken  and  brighter,  almost  cadmium-orange 
(R.  vi,  2)  anteriorly,  originating  behind  the  former  and  descending  on 
the  neck  to  past  the  middle;  a  third  band,  paler  buff,  curving  around 
the  corner  of  the  mouth  almost  meeting  below  the  corresponding  band 
of  the  other  side  on  the  posterior  third  of  the  neck;  a  fourth,  median 
band  between  the  last  ones;  soft  skin  of  feet  almost  "plumbeous" 
(R.  ii,  15)  underneath. 

Four  other  specimens  of  the  same  brood,  Nos.  61083-86,  are  colored 


38  bulletin:  museum  of  comparative  zoology 

essentially  as  the  above.  The  numbers,  sizes,  shape  and  arrangement 
of  the  dark  blotches  on  the  carapace  vary  to  a  great  extent  and  so  does 
consequently  the  light  network  (really  the  ground  color  of  the  cara- 
pace) separating  them,  it  being  slightly  wider  in  the  specimen  figured 
(pi.  19,  fig.  a).  There  is  one  feature  common  to  all  and  of  some  signifi- 
cance, viz.  the  more  or  less  parallel  arrangement  of  the  outer  blotches 
on  the  posterior  flap,  those  of  the  outer  row  nearly  forming  a  continu- 
ous dark  line,  the  next  row  also  coalescing  on  the  posterior  half,  thus 
clearly  indicating  the  blackish  submarginal  rings  of  the  other  species, 
as  shown  in  the  illustration  just  quoted. 

Wright  and  Funkliouser  (Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  67, 
1915,  p.  122)  note  that  "as  the  specimens  become  older,  the  gayly 
colored  markings  of  the  carapace  become  less  distinct  and  have  dis- 
appeared on  turtles  which  have  attained  a  length  of  6 inches  [152  mm.]." 
The  National  Museum  has  a  specimen  (No.  56804)  from  Irwin  Co., 
Georgia,  the  carapace  length  of  which  is  100  mm.,  which  still  shows  the 
pattern  as  described  above,  but  the  tubercles  on  the  disk  are  all  well 
developed. 

Their  further  statement  that  with  age  "the  plastron  grows  lighter  in 
color  and  the  head  uniformly  darker  with  the  markings  obsolete",  is 
borne  out  by  the  following  description  made  in  July  1919  of  a  live  male 
specimen  (now  U.S.N.M.  No.  62217)  from  Georgia,  Berrien  County, 
with  a  leathery  disk  about  235  mm.  long  and  195  mm.  wide:  Iris 
brownish  gray  with  a  brassy  edge  against  the  pupil.  Carapace  dark 
"raw  umber"  (R.  iii,  fig.  14)  with  large  dusky,  more  or  less  confluent 
irregular  marblings;  top  of  head,  neck  and  legs  "sepia"  (R.  iii,  fig.  3) 
with  paler  marblings ;  proboscis,  snout,  sides  of  head,  and  lips  strongly 
washed  with  "cinnamon"  (R.  iii,  fig.  20);  a  dark  brown  mark  across 
the  forehead  in  front  of  eyes  continued  behind  them  to  the  ear  as  a 
series  of  spots,  and  a  few  spots  of  the  same  color  behind  corner  of 
mouth;  on  underside  of  neck  back  of  the  skull  three  longitudinal  white, 
dusky -edged  marks;  underside  of  body  white;  digits  and  webs  strongly 
washed  with  plumbeous  (R.  ii,  fig.  15). 

Some  larger  specimens  may  still  retain  traces  of  the  original  pattern. 
A  full  grown  male  (U.S.N.M.  No.  60496,  carapace  length  316  mm.) 
collected  at  Auburndale,  Florida,  was  received  alive  on  February  28, 
1918,  and  its  colors  at  once  described  by  me  as  follows:  Iris  dark  silvery 
gray  with  a  darker  horizontal  bar  and  the  inner  edge  forming  a  com- 
plete bright  narrow  silvery  ring  sharply  defining  the  pupil;  inside  of 
nostrils  dark  pink;  inside  of  mouth  pale  flesh-color.  Ground  color  of 
carapace  bistre  with  anastomozing,  ill-defined  lines  (10  to  15  mm.  wide) 


stejneger:  American  soft-shell  turtles  39 

of  tawny  olive,  which  isolate  islands  of  the  ground  color  of  varying 
sizes  but  averaging  perhaps  25  mm.  in  diameter;  neck  and  legs  above 
dark  olive,  head  anteriorly  suffused  with  cinnamon  changing  to  pink 
on  proboscis;  underside  pinkish  white,  on  neck  suffused  with  dark 
purplish  gray  on  which  dusky-edged  cream-colored  lines  here  and  there 
with  touches  of  orange;  palms  and  soles  dark  olive. 

Aberrant  Color  Phase.  A  color  anomaly  shown  by  a  specimen,  the 
history  and  description  of  which,  because  of  the  uniqueness  of  the  pig- 
mentation deserves  to  be  recorded,  is  in  the  National  Museum. 

On  April  11,  1881,  there  was  entered  in  the  Museum  register  of 
reptiles  a  consignment  of  living  specimens  collected  in  March  by  James 
Bell  at  Gainesville,  Florida.  Among  them  was  an  "Aspidonectes"  of 
which  the  artist  Z.  Schindler  on  April  9  had  made  a  water-color  sketch 
for  use  in  later  painting  the  plaster  cast  for  the  "exhibition  series". 
The  specimen,  a  female  (Xo.  10545,  carapace  225  mm.  long,  16S  mm. 
wide),  now  a  skin  with  skull  separate,  the  color  sketch,  and  the  painted 
plaster  cast  (slightly  broken)  are  still  in  the  Museum.  The  water-color 
sketch  and  the  oil-painted  cast  agree  in  all  essentials,  except  that  the 
cast  is  (now)  considerably  darker.  The  skull  and  other  structural 
characters  are  those  of  a  normal  Amydaferox,  but  the  color  is  an  ex- 
treme case  of  erythrochroism.  The  ground  color  of  the  carapace  in 
the  water-color  agrees  closely  with  what  Ridgway  calls  "mummy 
brown"  (R.  iii,  fig.  10),  while  in  the  cast  it  is  more  like  his  "chestnut" 
(R.  iv,  fig.  9);  the  throat  and  front  of  neck  are  "rufous"  (R.  iv,  fig.  7) 
in  both,  though  lighter  in  the  former;  top  of  head,  back  of  neck,  and 
upper  side  of  legs  raw  umber  (R.  iii,  fig.  14)  in  the  water-color  sketch 
while  in  the  cast  they  are  dark  sepia  washed  with  rufous;  the  most 
conspicuous  feature  is  the  band  from  beneath  the  eye  on  the  auricular 
and  temporal  region  which  is  bright  vermilion  in  both  paintings- 
reminding  one  of  the  corresponding  band  in  Pseudemys  elegans)  and 
edged  with  a  narrow  dusky  line  anteriorly  above  and  below.  The  un- 
derside, judging  from  the  color  sketch  (and  the  specimen)  was  uniform 
white. 

Geographical  distribution 

Florida  north  to  South  Carolina  and  west  along  the  Gulf  Coast  to 
Louisiana. 


40 


bulletin:  museum  of  comparative  zoology 


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stejneger:  American  soft-shell  turtles 


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42 


bulletin:  museum  of  comparative  zoology 


List  of  specimens  in  U.  S.  National  Museum 


4373 

Fla. 

,  Palatka 

T.  Glover 

7651 

n 

ii 

(i 

8708 

Ga. 

Milledgeville 

T.  H.  Bean 

July,  1876 

8899  9  ad. 

it 

St.  Johns  R. 

S.  F.  Baird 

Apr.  3,  1877 

9670  d1  adol. 

S.  C 

!.,  Charleston 

C.  C.  Leslie 

May,  1878 

10545  9  adol. 

Fla. 

,  Gainesville 

J.  Bell 

Mar.,  1881 

10704 

it 

tt 

it 

19621  adol. 

Ga. 

Darien 

? 

20189  adol. 

Fla. 

,  Eustis 

T.  Holm 

March,  1893 

26035  juv. 

n 

ponds  near 

Welaka 

W.  C.  Kendall 

Mar.  20,  1897 

29210  9  ad. 

tt 

E.  A.  Mearns 

029339  adol. 

II 

Eustis 

H.  J.  Webber 

029448-9 

It 

Kissimmee 

E.  A.  Mearns 

029450-1 

029452-9  c?  ad. 

u 

n 

<( 

029460-2  c?  ad. 

11 

it 

a 

029463  tf1  ad. 

It 

a 

a 

029464-6  cf  ad. 

tt 

tt 

tt 

029467-8 

It 

it 

a 

029470  cf  ad. 

It 

a 

a 

029474  9  ad. 

tt 

it 

tt 

029475  9  ad. 

tt 

tt 

a 

029619  adol. 

Ga., 

Mimsville 

C.  S.  Brimley 

38123  9  ad. 

Fla. 

M.  L.  Odell 

38980-1  juv. 

Ga., 

Mimsville 

C.  S.  Brimley 

June  19,  1909 

51184  juv. 

Fla. 

ponds  near 

Tampa 

Evermann  &  Kendall 

Nov.    3,  1896 

51417-20  hf-gr. 

it 

St.  Petersburg 

C.  S.  Brimley 

51421  juv. 

tt 

Orlando 

it 

52476-83 

Fla., 

Eureka 

C.  S.  Brimley 

Aug.  12,  1915 

55316  c*  ad. 

Fla. 

Vero 

I.  M.  Weills 

56804  juv. 

Ga. 

Irwin  Co. 

J.  Hurter 

Oct.   15,  1902 

56805 

Fla., 

Orlando 

a 

July,  1911 

56806 

it 

Hillsboro  Co. 

tt 

June  15,  1910 

56807  ad. 

tt 

Bronson 

tt 

1891 

59318 

it 

Sebastian 

F.  Harper 

Feb.  23,  1917 

59727-8  (skullk 

only)  <?  ad. 

Fla., 

Lake  Okeechobee  F.  Harper 

Feb.    6,  1917 

60496  c?  ad. 

a 

[Auburndale] 

N.  R.  Wood 

Feb.,  1918 

60532  adol. 

a 

Auburndale 

a 

<( 

60533  ad. 

a 

(t 

it 

u 

60534  9  ad. 

a 

a 

a 

u 

stejneger:  americais 

r    SOFT-SHELL   TURTI 

,ES                         <i«5 

60547  &  ad. 

Fla.,  Auburndale 

N.  R.  Wood 

1918 

60556  juv. 

«                     a 

■  a 

Mar.,  1918 

60S28  eggs 

"     Homestead 

C.  A.  M osier 

60902-4 

"     Eureka 

C.  S.  Brimley 

1918 

61031 

"     Gulf  port 

A.  G.  Reynolds 

61083-7  pull. 

"     Auburndale 

N.  R.  Wood 

July  22,  1918 

61096-109 

ii                it 

u 

Aug.  24,  1918 

61352  9  ad. 

"     Lake  Miakka 

C.  M.  Barrett 

June  18,  1918 

62217  &  adol. 

Ga.,  Banks  Mill  Pond 

U.S.  Bur.  Fish. 

63343 

Fla.,  Auburndale 

N.  R.  Wood 

1920 

70398 

Ga.,  Mimsville 

C.  R.  Brimley 

71068-9 

? 

? 

71156  adol. 

Fla.,  Plant  City 

C.  R.  Aschemeier 

1926 

71681 ! 

S.C.,  Greenwood 

Dr.  Barrett 

73199 

Fla.,  Delray 

J.  D.  Thieme 

80963 

Fla. 

F.  H.  Benjamin 

Apr.,  1930 

84079 

"     15  mi.  from 

. 

Miami 

M.  K.  Brady 

1930 

84080 

"     Orlando 

E.  T.  Evans 

ii 

84603  pull. 

Ga.,  Chesser's  Id., 

Okefinokee  Swamp 

F.  Harper 

July  21,  1931 

86492  d* 

Fla.,  15  mi.  from 

Miami 

M.  K.  Brady 

Mar.,  1932 

86828  <?  adol. 

"     nr.  Birdon 

P.  Bartsch 

Aug.,  1932 

95767  <?  adol. 

"     Lake  Iamonia 

C.  R.  Aschemeier 

Feb.  16,  1935 

103736  Juv. 

"     Silver  Lake 

ti 

Mar.    7,  1938 

Amyda  spinifera  (Lesueur) 

Agassiz,  in  1857,  while  demonstrating  the  distinctness  of  Lesueur's 
species  described  30  years  before  from  the  Wabash  River  against  the 
contention  of  contemporary  authors  that  it  was  only  the  young  of 
Amyda  ferox,  at  the  same  time  established  three  additional  species, 
Aspidonectes  as  per  from  the  state  of  Mississippi,  A.  nuchalis  from 
Tennessee,  and  A.  emoryi  from  Texas.  The  latter  has  been  generally 
accepted,  nuchalis  has  been  generally  ignored,  and  asper  only  recently 
recognized  as  a  distinguishable  race  of  A.  spinifera.  As  such  it  will  be 
treated  below  trinominally. 

The  characters  attributed  to  A.  nuchalis  have  lost  their  significance 
after  the  accumulation  of  additional  material  of  A.  spinifera,  and  the 
study  of  available  series  of  specimens  in  connection  with  the  present 
investigation  have  not  yielded  data  indicating  a  separable  group  popu- 
lating the  upper  reaches  of  the  Tennessee  and  Cumberland  Rivers. 

'Identity  questioned. 


44  bulletin:  museum  of  compakative  zoology 

AMYDA  SPINIFERA  SPINIFERA1  (LeSUeUr) 

Plates  11-15 

1825. — Trionyx  ferox  SAY,  Journ.  Acad.  Nat.  Sci.,  ser.  1,  4,  pt.  2,  p.  218  (part: 
Mississippi,  Ohio,  and  Missouri  Rivers;  New  York,  etc.)  (not  of 
Schneider).— LeCONTE,  Ann.  Lync.  Nat.  Hist.,  New  York,  3,  1830, 
p.  93  (part:  all  streams  which  run  into  the  Mississippi). — HARLAN, 
Med.  Phys.  Res.,  1835,  p.  158  (part).— SCHLEGEL,  Fauna  Japon., 
Rept.,  1838,  p.  30,  pi.  5,  fig.  5  (Cumberland,  Tennessee  and  Ohio 
Rivers).— HOLBROOK,  North  Amer.  Herpet.,  1  ed.,  4,  1840,  p.  9 
(part);  2  ed.,  2,  1842,  p.  11,  pi.  1  (part:  Mississippi;  great  northern 
lakes;  Mohawk  River,  New  York). — DeKAY,  Zool.  New  York,  pt. 
3,  Rept.,  1842,  p.  6,  fig.  11  (Mohawk  River;  Hudson  River  near 
Albany,  New  York).— TROOST,  Seventh  Geol.  Tennessee,  1844, 
p.  39  (Tennessee).— THOMPSON,  Hist.  Vermont,  1853,  p.  29 
(Vermont:  Rivers  Lamoille  and  Winooski). — GRAY,  Cat.  Tort. 
Brit.  Mus.,  1844,  p.  49  (part);  Cat.  Shield  Rept.,  1,  1856,  p.  68 
(part).— KENNICOTT,  Trans.  Illinois  Agric.  Soc,  1,  1855,  p.  591 
(Illinois:  Cook  Co.,  Lake  Michigan).— STRAUCH,  Mem.  Acad. 
Sci.  St.  Petersbourg,  ser.  7,  5,  no.  7,  1862  (part);  8,  no.  13,  1865,  p. 
122  (part).— DITMARS,  Reptile  Book,  1907,  pi.  27,  upper  fig.  (not 
of  text).— GADOW,  Cambridge  Nat.  Hist.,  8,  1901,  p.  408  (part), 
p.  409,  fig.  92  (North  America). 
Amyda  ferox  ORTENBURGER,  Copeia,  no.  170,  May  1929,  p.  12 
(Oklahoma:  LeFlore  Co.);  p.  28  (Oklahoma:  Rogers  Co.). 

1827 .—Trionyx  spiniferus  LESUEUR,  Mem.  Mus.  Hist.  Nat.  Paris,  15,  Dec- 
1827,  p.  258,  pi.  6  (type-locality,  Wabash  River,  New  Harmony* 
Indiana;  types  in  Paris  Mus.;  Lesueur,  collector).  — WIED,  Reise 
Nord-Amerika,  1,  pt.  3,  1838,  pp.  140,  141  (Pittsburgh,  Pa.);  Voy. 
Amer.  Nord,  3,  1843,  p.  242.— HAY,  Indiana  Geol.  17  Rep.,  1892,  p. 
554;  Batr.  Rept.  Indiana,  1893,  p.  146  (Indiana:  generally  distri- 
buted).—HURTER,  Trans.  Acad.  Sci.  St.  Louis,  6,  1892,  p.  260 
(Missouri:  Mississippi  River,  Merimac  River,  and  Illinois  River). — 
HAHN,  Proc.  U.  S.  Nat.  Mus.,  35,  Dec.  1908,  p.  567  (Lawrence  Co., 
111.).— SIEBENROCK,  Zool.  Jahrb.  Suppl.,  10,  pt.  3,  1909,  p.  604 
(Mississippi  River  and  tributaries;  St.  Lawrence  River;  Hudson 
River) ;  Verh.  Zool.-Bot.  Ges.  Wien,  73,  Aug.  1923,  p.  186.— THOMP- 
SON, Thirteenth  Rep.  Michigan  Acad.  Sci.,  1911,  pp.  106,  107,  fig.  1 
(Cass  Co.,  Michigan).— ELLIS  and  HENDERSON,  Univ.  Colorado 
Stud.,  10,  no.  2,  May  1913,  p.  112  (Colorado:  Weld  Co.:  Evans, 
Cache  la  Poudre,  South  Platte  River,  and  Greeley). 

1  Spine  bearing,  with  reference  to  the  pointed  shape  of  the  tubercles  margining  the  anterior 
flap  of  the  leathery  carapace,  characteristic  of  older  specimens. 


stejneger:  American  soft-shell  turtles  45 

Gymnopus  spiniferus  DUMERIL  and  BIBRON,  Erpet.  Gen.,  2,  1835, 
p.  477  (atlas,  pi.  22,  fig.  1)  (Wabash  River).— DUMERIL,  Cat. 
Meth.  Coll.  Rept.  Mus.  Hist.  Nat.  Paris,  1851,  p.  22  (Wabash  River). 
— SAGER,  Peninsular  Journ.  Med.  Collat.  Sci.,  3,  no.  8,  1856,  p.  361 
(anatomy). — WIED,  Nova  Acta  Acad.  Leopold. -Carol.,  32,  pt.  1, 
1865,  p.  48  (Wabash  River). 

Gymnopodus  spiniferus  DUMERIL,  Arch.  Mus.  Hist.  Nat.,  7,  1856, 
p.  203. 

Aspidonectes  spiniferus  RHOADS,  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
1895,  p.  386  (Tennessee:  Samburg,  Obion  Co.). 

Aspidonectes  spinifer  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  1, 
1857,  p.  403;  vol.  2,  pi.  6,  figs.  1-2  (Lake  Champlain;  Lake  Ontario 
and  Erie;  New  York,  Pennsylvania,  Ohio,  Indiana,  Illinois,  Missouri, 
Michigan,  Wisconsin,  Iowa,  Ft.  Union,  Montana). — MILES,  First 
Biennial  Rep.  Geol.  Surv.  Michigan,  1861,  pp.  232,  233  (Michigan: 
as  far  W.  as  Genessee  Co.). — ALLEN,  Proc.  Boston  Soc.  Nat.  Hist., 
1874,   p.   69    (Musselshell   and  Yellowstone   Rivers,    Montana). — 
COPE,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  1875,  p.  51.— JORDAN,  Man. 
Vert.  North  Amer.,  ed.  3,  1880,  p.  168;  ed.  5,  1888,  p.  206  (Canada 
to   Kentucky   and   Minnesota);   ed.   8,    1899,   p.   206.— CRAGIN, 
Trans.  Kansas  Acad.  Sci.,  2,  1881,  p.  116  (Kansas:  Franklin  and 
Douglas  Cos.).— CONES  and  YARROW,  Bull.  U.  S.  Geol.  Geogr. 
Surv.,  4,  1878,  p.  261.— SMITH,  Rep.  Geol.  Survey  Ohio,  4,  1882, 
p.  668  (all  streams  flowing  into  Ohio  and  Lake  Erie). — TRUE,  Bull. 
U.  S.  Nat.  Mus.,  No.  24,  1883,  p.  29  (Webster  City,  Iowa;  Mt. 
Carmel,  Illinois;  Fox  River,  Illinois;  Ft.  Laramie,  Nebraska);  Fish. 
Industr.  United  States,  Sect.  1,  1884,  p.  152.— HOY,  Geol.  Wiscon- 
sin, Surv.  1873-1879,  1,  1883,  p.  423  (Western  Wisconsin).— DAVIS 
and  RICE,  Illinois  State  Lab.  Nat.  Hist.,  Bull.  No.  5,  1883,  p.  52 
(north  of  Ohio  River);  Bull.  Chicago  Acad.  Sci.,  1,  no.  3,  1883,  p.  32 
(Illinois).— HOY,  Geol.  Wisconsin,  1,  1883,  p.  423  (Wisconsin).- 
HUGHES,  Bull.   Brookville  Soc.   Nat.  Hist.,   No.  2,   1884,  p.  41 
(Franklin  Co.,  Indiana).— STOCKWELL,  Journ.  Comp.  Med.  Surg., 
9,  1888,  p.  28.— HIGLEY,  Trans.  Wisconsin  Acad.  Sci.,  7,  1889,  p. 
159    (south  and  west  Wisconsin). — GARMAN,   H.,   Bull.    Illinois 
State  Lab.  Nat.  Hist.,  3,  1892,  p.  246  (throughout  Illinois:  Rock 
Creek,  Kendall  Co.;  Piano;  Oregon,  Ogle  Co.;  Peoria,  Peoria  Co.; 
Bluff  Lake,  Union  Co.;  Wabash  River,  Mt.  Carmel).— KIRSCH, 
Bull.  U.  S.  Fish  Coram.  1894  (1895),  p.  81  (Eel  River,  Indiana).- 
McLAIN,  Notes  Coll.  Rept.  Arkansas,  1899,  p.  1   (Bloomington, 
Monroe   Co.,    Indiana). — SMITH,    Proc.    Linn.    Soc.    New   York, 
1898-99,  No.  11,  1899,  p.  24.— RAMSEY,  Proc.  Indiana  Acad.  Sci., 
1900  (1901),  p.  224  (Lake  Winona,  Indiana).— ATKINSON,  Ann. 
Carnegie  Mus.  Pittsburgh,  1,  1901,  p.   154   (Monongahela  River, 


46  bulletin:  museum  of  comparative  zoology 

Pennsylvania).— MORSE,  Ohio  Natural.,  1,  no.  8,  1901,  p.  127 
(Columbus,  Ohio);  Proc.  Ohio  Acad.  Sci.,  4,  pt.  3,  Spec.  Pap.  No.  9, 
1904,  p.  138  (Ohio:  Columbus;  Sandusky;  London).— PAULMIER, 
New  York  State  Mus.  Bull.  51,  1902,  p.  392  (Lakes  Ontario  and 
Erie;  through  Erie  Canal  to  Hudson  River). — CLARK,  Rep.  4 
Michigan  Acad.  Sci.,  1904,  p.  193  (Eaton  Co.,  Michigan).— JOR- 
DAN, Man.  Vert.  Anim.  U.  S.,  1904,  p.  206.— GIBBS,  NOTE- 
STEIN  and  CLARK,  Rep.  7,  Michigan  Acad.  Sci.,  1905,  p. 
110  (Michigan:  Brookfield,  Ann  Arbor,  Olivet,  Kalamazoo,  Van 
Buren,  Montcalm  and  Allegan  Cos.). — STONE,  Amer.  Natural., 
40,  1906,  p.  168  (into  Delaware  Valley;  Cooper's  Creek  and  Warren 
Co.,  New  Jersey;  Allegheny  River,  western  Pennsylvania). — NASH, 
Check  List  Vertebr.  Ontario,  1906,  p.  17  (Canada:  Ontario,  western 
part;  one  record  Ottawa  River). — FOWLER,  Rep.  New  Jersey  State 
Mus.,  1906  (1907),  p.  211,  pi.  57  (Cooper's  Creek;  Paulin's  Kill, 
Hainesburg,  Warren  Co.,  New  Jersey,  introduced). — SURFACE, 
Zool.  Bull.  Pennsylvania  Dep.  Agric,  6,  nos.  4-5,  1  Sept.,  1908,  p. 
121,  fig.  2  (Pennsylvania:  Indiana  Co.;  Somerset  Co.). — REED  and 
WRIGHT,  Proc.  Amer.  Philos.  Soc,  48,  1909,  p.  408  (Cayuga  Lake, 
New  York). 
Trionyx  spinifer  BOULENGER,  Cat.  Chel.  Brit.  Mus.,  1889,  p.  259 
(Foxbury,  Pennsylvania;  Wabash  River). — SCHNEE,  Zeitschr. 
Naturwiss.,  72,  pt.  3,  Dec.  24,  1899,  p.  197.— SIEBENROCK,  Sitz. 
Ber.  Akad.  Wiss.  Wien,  Math.-Nat,  KL,  111,  1902,  p.  829,  fig.  10 
(plastron).— HENSH AW,  Occas.  Pap.  Boston  Soc.  Nat.  His.,  7, 
pi.  1,  1904,  p.  4  (Lake  Champlain,  Vermont).— DITMARS,  Rep- 
tile Book,  1907,  p.  77,  pi.  26,  upper  and  middle  figs.;  pi.  28  (Quincy, 
Adams  Co.,  Illinois). 

Platypeltis  spinifer  BAUR,  Proc.  Amer.  Philos.  Soc,  31,  1893,  p.  220 
(Wabash  River,  Indiana).— EVERM ANN  and  CLARK,  Proc. 
Indiana  Acad.  Sci.,  1916  (1918),  p.  473  (Lake  Maxinkuckee,  Indiana). 

Platyrettis  KIRSCH,  Bull.  U.  S.  Fish  Comm.  1894  (1895),  p.  333 
(Maumee  River  Basin,  Ohio  and  Indiana). 

Amyda  spinifer -POTTER,  Copeia,  No.  82,  Oct.  15,  1921,  p.  76  (Thorn- 
apple  River,  Barry  Co.,  Michigan).— SOLA,  Bull.  New  York  Zool. 
Soc,  34,  no.  5,  Sept.-Oct,,  1931,  pp.  134,  141,  155,  fig.  6  (Lake  Cham- 
plain). 

Callinia  spinifera  GRAY,  Proc  Zool.  Soc.  London,  1869,  p.  222 
{spicifera  err.  typogr.);  1873,  (p.  60,  figs.);  Suppl.  Cat.  Shield.  Rept. 
Brit,  Mus.,  pt.  1,  1870,  p.  109. 

Amyda  spinifera  HURTER,  Trans.  Acad.  Sci.  St.  Louis,  20,  1911,  p. 
251  (Missouri:  Mississippi,  Missouri,  Osage,  Gasconade,  Meramee 
and  White  Rivers).— STEJNEGER  and  BARBOUR,  Check  List 


stejneger:  American  soft-shell  turtles  47 

North  Amer.  Amph.  Rept.,  Ed.  1,  Dec.  12,  1917,  p.  125  (Mississippi 
River  and  tributaries  west  to  Colorado,  north  to  Montana;  St. 
Lawrence  River  and  tributaries;  east  to  Vermont,   western  New 
York  and  Pennsylvania);  Ed.  2,  1923,  p.  141;  Ed.  3.,  1933,  p.  153.- 
WRIGHT,  Copeia,  No.  66,  Feb.  25,  1919,  p.  8  (Bays  of  Lake  On- 
tario, New  York).— BABCOCK,  Mem.  Boston  Soc.  Nat.  Hist.,  8, 
no.  3,  April  1919,  p.  419  (east  shore,  Lake  Champlain,  Vermont). — 
CLARK  and  SOUTHALL,  Rep.  U.  S.  Comm.  Fish  1919  (1920), 
App.  no.  7,  p.  15,  pis.  7-8  (economics).— EVERMANN  and  CLARK, 
Lake    Maxinkuckee,    1920,    p.    592    (habits).— ORTENBURGER, 
Copeia,  No.  99,  Oct.  15,  1921,  p.  76  (Decatur  Co.,  Indiana);  Proc. 
Oklahoma  Acad.  Sci.,  6,  pt.  1,  1926,  p.  100  (McCurtrin  and  Push- 
nataha  Cos.,  Oklahoma). — BLANCHARD,  Occas.  Pap.  Zool.  Mus. 
Univ.  Michigan,  No.  117,  July  6,  1922,  p.  18  (Reelfoot  Lake,  West. 
Tennessee);  Univ.  Iowa  Studies  Nat.  Hist.,  12,  no.  2,  1923,  p.  24 
(Little  Sioux  Riv.,  Dickinson  Co.,  Iowa);  Pap.  Michigan  Acad.  Sci., 
5,    1925,   p.   386    (10  miles  W  of  Columbus,    Indiana).— WEED, 
Copeia,  No.   116,   March  15,   1923,  p.  48  (Meredosia,  Illinois).— 
BISHOP,  Copeia,  No.  125,  Dec.  31,  1923,  p.  120  (Albany  Co.,  New 
York).— PRATT,   Man.  Vert.  United  States,   1923,  p.  249.— LO- 
GIER,  Canad.  Field  Natural.,  39,  May,  1925,  p.  95  (Point  Pel6e; 
Hamilton,  Dundas  Marsh,  Ontario,  Canada);  Roy.  Ontario  Zool. 
Mus.  Handb.  no.  4,  1939,  p.  56,  pi.  7,  fig.  2;  pi.  8,  fig.  7  (Ontario: 
Hamilton  Bay  and  Dundas  Marsh,  Wentworth  Co.;  Thames  River 
at  Beachville,  Oxford  Co.;  Grand  River  at  Dunnville,  Haldimand 
Co.;   Long   Point,    Norfolk  Co.,   and  Point  Pelee,   Essex  Co.). — 
SCHMIDT,  Copeia,  no.  154,  May  20,  1926,  p.  132  (Chippewa  Co., 
Wise.).— MYERS,  Proc.  Indiana  Acad.  Sci.,  35,  1926,  p.  292  (In- 
diana); vol.  36,  1927,  p.  339  (Helmsburg,  Indiana). — BURT,  Occas. 
Pap.  Zool.  Mus.  Univ.  Michigan,  no.  189,  Dec.  12,  1927,  p.  9  (Riley 
Co.,  Kansas).— LINSDALE,  Copeia,  no.  164,  July-Sep.,  1927,  p.  81 
(Doniphan  Co.,  Kansas);  Trans.  Kansas  Acad.  Sci.,  36,  1933,  p.  208 
(Cowley  and  Lane  Cos.,  Kansas).— RUTH VEN,  THOMPSON,  and 
GAGE,  Herpet.  Michigan,  1928,  p.  163,  pi.  19,  fig.  3  (Michigan).- 
MERTENS,  Zool.  Gart.  Leipzig,  new  series,  1,  pt.  5-6,  1928,  p.  199 
(distr.  New  York  State  by  canals). — GLOYD,  Trans.  Kansas  Acad. 
Sci.,  31,  1928,  p.  135  (Franklin  Co.,  Kansas).— POPE  and  DICKIN- 
SON, Bull.  Publ.  Mus.  Milwaukee,  8,  no.  1,  Apr.  3,  1928,  p.  82,  pi.  21, 
figs.    7-8    (Wisconsin:   Burnett,    Crawford,    Grand,   Oneida,    Polk, 
Waukesha,  Chippewa,  Washburn,  and  Pepin  Cos.). — POPE,  Year 
Book  Mus.   Milwaukee,   1928   (1929),  pp.    180,   183   (Vernon  and 
LaCrosse  Cos.,  Wisconsin). — JORDAN,  Man.  Vert.  Northeast  U.S. 
(13  ed.)  1929,  p.  254.— CAHN,  Copeia,  no.  170,  May  1929,  p.  8 
(Lake   La   Belle,    Waukesha   Co.,    Wisconsin). — FORCE,    Copeia, 
1930,  no.  2,  p.  38  (Tulsa  Co.,  Oklahoma).— DOLLEY,  Amer.  Mid- 


48  bulletin:  museum  of  comparative  zoology 

land  Natural.,  14,  no.  3,  May,  1933,  p.  203  (St.  Joseph  Riv.,  Berrien 
Co.,  Michigan).— RUST,  Blatt.  Aquar.  Terrarienk.,  45,  1934,  p.  — , 
sep.  p.  12.— BURT  and  HOYLE,  Trans.  Kansas  Acad.  Sci.,  37, 

1934,  p.  198  (Kansas).— BURT,  Amer.  Midland  Natural.,  16,  no.  3, 

1935,  p.  321  (Kansas:  Barber,  Reno  and  Sedgwick  Cos.).— BOYER 
and  HEINZE,  Trans.  Acad.  Sci.  St.  Louis,  26,  no.  4,  Apr.  1,  1934,  p. 
199  (Missouri:  Jefferson  Co.,  common).— NETTING,  Nawakwa 
Fireside  (N.S.),  nos.  3-4,  Apr.,  1935,  p.  49  (Penna,:  8  counties); 
Proc.  Pennsylvania  Acad.  Sci.,  10,  1936,  p.  27  (Pennsylvania,  In- 
diana Co.,  Plum  Creek  and  Crooked  Creek).— BRINN,  Rep.  Penn- 
sylvania Fish  Comm.  for  1938-1939,  p.  127  (Pennsylvania:  rec.  from 
9  counties  in  Ohio  and  Lake  Erie  drainages). — SCHMIDT  and 
NECKER,  Bull.  Chicago  Acad.  Sci.,  5,  no.  4,  Sept.  27,  1935,  p.  76 
(Illinois:  Cook  Co.,  Kankakee  Co.;  Indiana:  Lake  Co.).— TAYLOR 
Univ.  Kansas  Sci.  Bull.,  22,  no.  11,  Apr.  15,  1935,  p.  217  (Arkansas: 
Devall  Bluff,  Prairie  Co.;  Lewisville,  Lafayette  Co.).— BABBITT, 
Bull.  Boston  Soc.  Nat.  Hist.,  no.  78,  Jan.  1936,  p.  10  (Lake  Cham- 
plain,  Vermont,  rare). — HIBBARD,  Trans.  Kansas  Acad.  Sci.,  39, 

1936,  p.  281  (Mammoth  Cave  Nat.  Park,  Kentucky).— PARKER, 
Jour.  Tennessee  Acad.  Sci.,  12,  no.  1,  Jan.,  1937,  p.  85,  fig.  18  (Bayou 
du  Chien;  Reelfoot  Lake,  Tennessee);  Rep.  Reelfoot  Lake  Biol. 
Sta.,  3,  Jan.  1939,  p.  88  (Tennessee:  Reelfoot  Lake).— CAHN, 
Illinois  Biol.  Monogr.  16,  Aug.  31,  1937,  p.  184,  pis.  25-27,  30  fig.  b, 
map  20   (Illinois).— GREETE,  Herpetologica,  1,  no.  4,  Nov.   16, 

1937,  p.  116  (West  Virginia:  Randolph  Co.,  Tygart  River  near  El- 
kins). — ALEXANDRE,  Soc.  Canadienne  Hist.  Nat.,  tract  no.  39, 
Apr.  1937,  p.  2  (Canada:  Quebec:  Lake  Champlain  and  Richelieu 
River,  tributary  to  St.  Lawrence  R.,  right  side).— HENNING, 
Copeia,  1938,  no.  2,  June  30,  1938,  p.  92  (Boone  Co.,  Missouri).— 
DENNING  and  BLACK,  Occas.  Pap.  Univ.  Arkansas  Mus.,  No.  1, 
June  1938,  p.  46  (Arkansas:  Lafayette,  Prairie,  Chicot,  Clay,  Gar- 
land, Lawrence,  and  Washington  Cos.). — CONANT,  Amer.  Mid- 
land Natural.,  20,  no.  1,  1938,  p.  157,  pi.  21,  fig.  1  (right),  pi.  22, 
figs.  1,  2  (Ohio:  map);  Herpetologica,  1,  no.  5,  Dec.  30,  1938,  p.  138 
(Ohio:  Lucas  Co.).— NECKER,  Bull.  Chicago  Acad.  Sci.,  6,  no.  1, 
1939,  p.  10  (Illinois:  Cook  Co.:  Evanston;  Kankakee  Co.:  Kankakee 
Riv.  near  Altort).— WELTER  and  CARR,  Copeia,  1939,  no.  3, 
Sept.  9,  p.  130  (Kentucky:  Triplet  Co.;  Fox,  Fleming  Co.;  rare  in 
East  Ky.).— LOGIER,  Canad.  Field-Nat.,  39,  May,  1925,  p.  95 
(Point  Pelee;  Hamilton,  Dundas  Marsh,  Ontario,  Canada);  Roy. 
Ontario  Zool.  Mus.  Handb.  no.  4,  1939,  p.  56,  pi.  7,  fig.  2;  pi.  8,  fig.  7 
(Ontario:  Hamilton  Bay  and  Dundas  Marsh,  Wentworth  Co.; 
Thames  River  at  Beachville,  Oxford  Co. ;  Grand  River  at  Dunnville, 
Haldimand  Co.;  Long  Point,  Norfolk  Co.;  and  Point  Pelee,  Essex 
Co.).— GENTRY,  Rep.  Reelfoot  Lake  Biol.  Sta,,  5,  Jan.  1941,  p. 


stejneger:  American  soft-shell  turtles  49 

75  (Tennessee:  Clay  Co.,  Overton  Co.);  Journ.  Tennessee  Acad. 
Sci.,  16,  no.  3,  1941,  p.  332  (Tennessee:  Overton,  Fentress,  Pickett, 
Jackson,  and  Clay  Cos.).— CAGLE,  Copeia,  1942,  no.  3,  Oct.  8, 
p.  155  (Illinois:  Jackson  and  Williamson  Cos.) 
Platypeltis  spinifera  RUTHVEN  and  THOMPSON,  Herpet.  Michi- 
gan, 1912,  p.  129  (Michigan).— THOMPSON,  Misc.  Pap.  Zool. 
Michigan,  1916,  p.  63  (Monroe  Co.,  Michigan). 

1827.— Trionyx  ocellatus  LESUEUR,  Mem.  Mus.  Hist.  Nat.  Paris,  15,  Dec. 
1827,  p.  261  (type-locality,  New  Harmony,  Indiana).— DeKAY, 
Zool.  New  York,  pt.  3,  Reptiles,  1842,  p.  7  (Mohawk  River,  New 
York). 

1838. — Trionyx  annulifer  WIED,  Reise  Nord-Amerika,  1,  pt.  3,  p.  140  (type- 
locality,  Ohio  River  at  Pittsburgh,  Pennsylvania);  Voy.  Amer. 
Nord.,  3,  1843,  pp.  242-243. 

1844.— Tyrse  argus  GRAY,  Cat.  Tort.  Brit.  Mus.,  p.  48  (type-locality,  "West 
Africa,  Sierra  Leone?";  type  in  Brit.  Mus.;  Lord  Derby,  collector); 
Knowlsley  Menag.,  1846  (pi.  — ). 
Trionyx  argus  GRAY,  Cat.  Shield  Rept.  Brit.  Mus.,  pt.  1,  March  8, 
1856,  p.  68. 

1856.  Trionyx  annularis  "Wied",  GRAY,  Cat.  Shield  Rept.  Brit.  Mus.,  pt. 

1,  March  8,  1856,  p.  69  (lapsus  in  synonymy). 

1857.  Aspidonectcs  nuchalis  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  1, 

p.  406  (type-localities,  Cumberland  and  Tennessee  rivers;  cotypes, 
Mus.  Comp.  Zool.,  nos.  1623-1625,  Cumberland  river).— COPE, 
Bull.  U.  S.  Nat.  Mus.,  no.  1,  1871,  p.  51.— TRUE,  Bull.  U.  S.  Nat. 
Mus.,  no.  24,  1883,  p.  5;  Fish.  Industr.  United  States,  sect.  1,  1884, 
p.  152  (middle  western  states).— DAVIS  and  RICE,  Illinois  State 
Lab.  Nat.  Hist.,  Bull.  No.  5,  1883,  p.  52  (Cumberland  and  Upper 
Tennessee  Rivers).— JORDAN,  Man.  Vert.  Anim.  North  United 
States,  ed.  5,  1888,  p.  206  (Cumberland  and  Upper  Tennessee  Rivers) ; 
ed.  8,  1899,  p.  206.— RHOADS,  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
1895,  p.  404  (Cumberland  River,  near  Nashville). 

1861.  Amyda  mutica  MILES,  First  Biennial  Rep.  Geol.  Surv.  Michigan,  pp. 
2323,  233  (Michigan).  (Not  of  Lesueur).— SMITH,  Science  News 
Suppl.,  1,  1879,  p.  7  (Michigan).— KIRSCH,  Bull.  U.  S.  Fish  Comm. 
1894  (1895),  p.  41  (Indiana:  Eel  River);  p.  333  (Maumee  Riv., 
Ohio). 

1865. — Gymnopus  olivaceus  WIED,  Nova  Acta  Acad.  Leopold.-Carol.,  32, 
pt.  1,  p.  55,  pi.  5  (type-locality,  New  Harmony,  Wabash  River, 
Illinois). 

1869.— Callinia  spicifera  GRAY,  Proc.  Zool.  Soc.  London,  1869,  pp.  222 
(lapsus  for  spinifera). 


50  bulletin:  museum  of  comparative  zoology 

1939.—  Amyda  spinifera  spinifera  STEJNEGER  and  BARBOUR,  Check 
List  N.  A.  Amph.  Rept.,  ed.  4,  p.  172;  ed.  5, 1943,  p.  213.— ANDER- 
SON, Bull.  Chicago  Acad.  Sci.,  6,  no.  11,  July  8,  1942,  p.  219  (Mis- 
souri: Jackson  Co.;  Missouri  River  near  Atherton). — EDGREN, 
Copeia,  1942,  no.  3,  Oct.  8,  p.  180  (Michigan:  Van  Buren  Co.,  Rey- 
nold Lake).— PETERS,  Copeia,  1942,  no.  3,  Oct.  8,  p.  183  (Illinois: 
Cumberland  Co.). 

Geographical  distribution 

Amyda  spinifera  has  been  credited  to  the  fauna  of  South  Carolina 
on  the  strength  of  the  specimen  U.S.N.M.  No.  7650  recorded  in  U.  S. 
Nat.  Mus.  Bulletin  No.  24  (1883),  p.  29,  as  coming  from  "Abbeville, 
S.  C".  It  is  also  the  record  upon  which  is  based  True's  record  of  the 
occurences  of  Aspidonectes  spinifer  in  H.  S.  Thompson's,  South  Caro- 
lina, 1883,  p.  238.  The  record  is  undoubtedly  erroneous,  the  ascertain- 
able facts  being  as  follows : 

The  specimen,  recently  hatched,  was  one  of  the  numerous  neglected 
turtles  found  in  the  collection  when  Dr.  G.  Brown  Goode  took  charge 
and  tintagged  and  registered  the  specimens.  He  entered  the  present 
one  in  1872  as  No.  7650,  the  tin  tag  bearing  that  number.  In  the  record 
book  [in  Brown  Goode's  handwriting]  it  is  noted  as  an  Aspidonectes 
[no  specific  name] ;  locality :  "Abbeville  S.  C";  and  no  further  remarks. 
There  is  now  no  other  indication  of  its  origin  than  an  old  torn  scrap 
of  paper  in  the  bottle  with  "7650  Abbeville  S.  C."  likewise  in  his  hand- 
writing. On  the  back  of  this  label,  however,  there  is — in  very  faded 
ink  and  in  an  entirely  different  handwriting — the  remnant  of  an  in- 
scription, beginning  and  end  clipped  off,  "ville  Mis".  If  this  remnant 
really  is  part  of  the  original  locality  record,  it  seems  probable  that  it 
may  have  read  "Abbeville,  Miss."  and  inadvertently  attributed  to 
South  Carolina  as  the  more  familiar  locality.  The  specimen  is  unques- 
tionably an  Amyda  spinifera.  The  whole  pattern  of  coloration  is 
normal  of  that  species,  which  of  course  at  once  excludes  A.  ferox.  The 
single  regularly  defined  submarginal  dusky  ring  on  the  upper  side  of 
the  disc  likewise  excludes  A.  agassizii,  the  species  one  would  expect 
if  it  were  collected  in  Abbeville,  S.  C.  To  make  perfectly  sure  of  its 
identity  I  have  had  the  skull  extracted.  It  shows  all  the  characteristic 
features  of  A.  spinifera  as  distinguished  from  A.  agassizii. 

Another  erroneous  record  of  Trionyx  spiniferus,  viz.  in  Pope's 
Turtles  (1931)  pi.  45,  figs.  98,  99)  showing  excellent  figures  of  two  soft- 
shelled  turtles  from  the  region  of  Columbia,  South  Carolina,  needs 
correction.   Both  pictures  are  plainly  of  Amyda  agassizii. 


stejneger:  American  soft-shell  turtles 


51 


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59263 

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o 
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70397 
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52  bulletin:  museum  of  comparative  zoology 

Stockwell's  reference  to  the  occurrence  of  Aspidonectes  spinifer 
"North  of  Athabasca  Lake"  (Journ.  Compar.  Medic.  Surg.,  9,  1888, 
p.  28)  must  rest  on  some  curious  lapsus. 

Where  Amy  da  spinifer  meets  with  A.  asper  is  as  yet  conjectural. 
It  extends  at  least  as  far  south  as  the  northern  part  of  the  State  of 
Mississippi,  for  we  have  undoubted  specimens  from  De  Soto  County 
(U.S.N.M.  No.  92606)  and  Lake  Washington,  Washington  County 
(No.  92607).  A  young  male  specimen  from  Madison  Co.,  Northern 
Louisiana  (No.  83985)  is  likewise  this  form  with  only  one  marginal 
stripe. 

The  Museum  of  Comparative  Zoology  has  a  specimen  from  Colum- 
bus, Ga  (M.C.Z.  1606). 

Mississippi  River  and  tributaries,  west  to  Colorado,  north  to  Mon- 
tana; St.  Lawrence  River  and  tributaries;  east  to  Vermont,  western 
New  York,  and  Pennsylvania. 

A.  nuchalis,  judging  from  the  slight  material  at  hand,  is  not  a 
strongly  differentiated  form.  The  character  chiefly  relied  on  by  Agas- 
siz,  "the  most  prominent  specific  character  .  .  .  the  marked  depression 
on  either  side  of  the  blunt  median  keel,"  does  not  hold  in  a  series  of 
specimens.  I  find  quite  a  number  so  characterized  among  typical  A. 
spinifer  a,  and  while  one  of  the  specimens  from  near  Sevier  ville,  Tenn., 
has  a  rather  flat  bony  carapace,  the  other  has  "the  blunt  keel,  which 
extends  along  the  median  line  and  slopes  uniformly  upon  the  sides," 
exactly  as  he  describes  it  diagnostically  for  A.  spinifera  (p.  404). 
The  angle  of  the  entoplastron,  in  the  few  examples  examined,  is  some- 
what more  obtuse,  between  90°  and  95°,  than  in  corresponding  speci- 
mens of  typical  A.  spinifera.  The  sharply  defined  ocelli  on  the  carapace 
seem  to  be  larger  than  in  A.  spinifera  of  the  same  age  (size). 

I  do  not  at  all  understand  Agassiz's  note  that  "this  species  differs 
strikingly  from  Asp.  spinifer  in  the  much  more  elongated  form  of  the 
male,  and  in  the  great  development  of  the  marginal  spines  and  of  the 
tubercles  upon  the  carapace,  which  project  very  slightly  in  the  male 
Asp.  spinifer."  On  the  contrary,  in  our  upper  Tennessee  specimens, 
presumably  typical  A.  nuchalis,  the  carapace  of  the  males  is  wider 
than  in  the  females,  and  the  spines  on  the  anterior  edge  very  much 
smaller,  exactly  as  in  typical  A.  spinifera. 

U.S.N.M.  86677  from  Cumberland  Gap,  and  86682  from  2  miles 
west  of  Sevierville,  Tennesse,  are  typical  "nuchales." 


stejneger:  American  soft-shell  turtles 


53 


List  of  specimens  in  the  U.  S.  National  Museum 


58 

Mont.,  Ft.  Union 

F.  V.  Hayden 

7163(029528) 

Tenn.,  Nashville 

J.  Varden 

7165(029529) 

it            it 

" 

7166(029530) 

it            a 

tt 

7167 

n            it 

tt 

7169  juv. 

tt            tt 

a 

7648  &  adol. 

Wyo.,  Ft.  Laramie 

F.  V.  Hayden 

7649 

? 

? 

7650  pull. 

Miss.,  Abbeville? 

? 

7661  juv. 

111. 

R.  Kennicott 

8359 

Ind.,  Madison  ? 

? 

9654 

111.,  Mt.  Carmel 

R.  Ridgway 

May, 

1878 

9717 

111.,  Mt.  Carmel 

Mrs.  L.  M.  Turner 

June, 

1878 

9928  juv. 

Iowa,  Webster  City 

C.  Aldrich 

1878 

11625  juv. 

? 

? 

11631  juv. 

La.,  Prairie  Mer  Rouge 

? 

12061  juv. 

111.,  Mt.  Carmel 

L.  M.  Turner 

14535  juv. 

Mont.,  Ft.  Custer 

C.  Bendire 

14536  9  ad. 

it              a 

tt 

16704  9 

Ala.,  Courtland 

rP.  H.  Kirch,  E.  O. 

■ 

Jones,  and 

May, 

1889 

16705 

ti          a 

W.  M.  Andrews 

tt 

17823  juv. 

Ark.,  Benton 

Jordan  &  Gilbert 

19622-3   9  &  adol. 

? 

19625 

? 

? 

21128-9  juv. 

Ohio,  Cuyahoga  River 

A.  J.  Woolman 

July 

25,  1893 

21416-7 

? 

? 

21567-8 

Ohio,  Edgerton 

P.  H.  Kirsch 

July 

28,  1893 

21569-70 

Ind.,  Fish  Cr.,  near 

Hamilton 

tt 

July 

21,  1893 

21571-7 

Ohio,  Maumee  Basin 

a 

1893 

22711 

Ind.,  Vincennes 

R.  Ridgway 

24536 

Mont.,  Ft.  Custer 

C.  Bendire 

Mar. 

8,  1886 

26290 

Ohio,  Franklin  Co. 

R.  C.  Osborn  &  E. 

C.  Williamson 

June, 

1897 

029014   9 

"     Columbus 

O.  Davie 

33494 

Ind.,  Lake  Maxin- 

kuckee 

B.  W.  Evermann 

July 

21,  1900 

33495 

Yellow  River  nor( 

h 

of  Burr  Oak 

tt 

Oct. 

3,  1900 

33496-501 

"      L.  Maxinkuckee 

1899- 

-1900 

33767 

W.  Va.,  Dry  Fork, 

Perryville 

W.  P.  Hay 

1900 

34               bulletin:  museum  of  comparative  zoology 

35404-8 

Ind.,  L.  Maxinkuckee 

H.  W.  Clark 

1900 

36412 

111.,  Illinois  R. 

S.  P.  Bartlett 

42583  juv. 

Ind.,  Long  Point,  L. 

Maxinkuckee 

Evermann  &  Clark 

Oct. 

5,  1906 

42584  juv. 

"      L.  Maxinkuckee 

n                a 

42585  eggs 

u                    a 

a                it 

Nov. 

17,  1906 

42905-6  juv. 

"       Burlington 

B.  W.  Evermann 

1899 

50670  juv. 

Twin  Lakes 

H.  W.  Clark 

July 

2,  1909 

51213  hf-gr. 

Mich.,  Monroe  Piers 

C.  Rutter 

Aug. 

13,  1894 

51214      " 

Ohio,  Toledo,  Grassy 

Point 

<< 

Aug. 

3,  1894 

51529  ad. 

Kansas 

R.  L.  Moodie 

53521 

Iowa,  Fairport 

Bur.  Fisheries 

Apr. 

24,  1911 

53522 

111.,  Hamilton 

J.  McAdams 

May 

1,  1915 

53523-6 

Iowa,  Fairport 

E.  Snyder 

Oct. 

1,  1914 

54421  cf  ad. 

Mont.,  Crow  Agency 

M.  A.  Hanna 

Aug. 

5,  1916 

54422-3 

it              a 

R.  Kellogg 

July 

8,  1916 

54730  9  adol. 

Iowa,  Fairport 

J.  Snyder 

June, 

1916 

54731    9  adol. 

a             tt 

a 

July 

2,  1916 

54732 

a             a 

tt 

a 

54739-41 

n             a 

a 

May 

8,  1916 

54743-6 

a            a 

ti 

May, 

1916 

54747 

111.,  Meredosia 

Freeland  &  Williams  1908 

55680 

"  Madison  Co. 

J.  Hurter 

55681 

"  Union  Co. 

!( 

55682 

Ky.,  Morgan  Co. 

It 

'55683 

Kans.,  Greenwood  Co. 

It 

July-Aug.,  1912 

55684 

Mo.  Stone  Co. 

It 

June  27,  1908 

55085 

"  St.  Louis  Co. 

It 

1913 

55686 

"  St.  Louis 

It 

55687 

it             a 

tt 

Apr. 

9,  1905 

55688 

"  Reynolds  Co. 

a 

July 

21,  1911 

55689 

"  FranklynCo. 

a 

Aug. 

20,  1911 

55690 

"  Washington  Co. 

tt 

June 

25,  1911 

59046 

"  St.  Louis 

ti 

May 

20,  1908 

59263-6  9  adol 

.  Miss.,  Homer 

F.  Schrader 

59267  9 

Mo.,  Alexandria 

E.  Stringham 

June 

5,  1916 

59270-5 

Minn.,  Homer 

F.  Schrader 

Sept. 

8,  1916 

59277 

111.,  between  Warsaw 

and  Hamilton 

E.  Stringham 

Aug. 

25,  1916 

59279-80 

Mo.,  Canton 

Bur.  Fisheries 

59285 

Iowa  ? 

n 

59736 

Mont.,  Crow  Agency 

R.  Kellogg 

July 

23,  1916 

59956 

Ind.,  Madison 

O.  P.  Hay 

59979  d"  juv. 

Ind.  ? 

ti 

stejneger:  American  soft-shell  turtles 


55 


60571  juv. 

III.,  Madison  Co. 

J.  Hurter 

70397  adol. 

Okla.,  Red  River, 

McCurtain  Co. 

A.  I.  Ortenburger 

1924 

72387 

Ind.,  Knox 

H.  W.  Clark 

Aug. 

12,  1909 

73668-9  juv. 

Miss.,  Greenwood 

I.  L.  Towers 

1925 

83985  <?  adol. 

La.,  2  mi.  E.  of  Mounds  C.  E.  Burt 

June 

30,  1931 

86677  9  ad. 

Tenn.,  5  mi.  S.E. 

Cumberland  Gap 

a 

July 

22,  1932 

86681-2  d"  adol 

..       "      2  mi.  W.  Sevierville 

1932 

87164  juv. 

"      2  mi.  S.  Kingston          " 

July 

27,  1932 

90441-4  juv. 

Kans.,  1  mi.  W.  Winfield 

Apr. 

30,  1933 

91022  juv. 

"      Winfield 

L.  Hoyle 

May 

28,  1933 

92606  pull. 

Miss.,  Lake  Cormorant  S.  F.  Hildebrand 

June 

9,  1933 

92607  juv. 

"     Greenville 

a 

May 

31,  1933 

93089-94  juv. 

Mo.,  Dardenne  Cr., 

St.  Charles  Co. 

L.  Hubricht 

Aug. 

16,  1931 

95140  pull. 

Miss.,  1  mi.  W.  Yazoo 

City 

C.  E.  Young 

Aug. 

13,  1934 

95261    9  ad. 

Kans.,  2  mi.  E.  of 

Calista 

C.  E.  Burt 

May 

25,  1934 

95301 

"      11  mi.  S.E.  of 

Winfield 

<< 

Aug. 

31,  1934 

95352   9  ad. 

Ark.,  7  mi.  N.W. 

Natural  Dam 

it 

June 

7,  1934 

95405  &  ad. 

Mo.,  Glaize  Creek, 

Jefferson  Co. 

A.  A.  Heinze 

July 

20,  1932 

99862-75 

La.,  Red  River  near 

Shaw 

C.  E.  Burt 

1935 

100160   9  ad. 

"    Bayou  Chene 

it 

June 

17,  1935 

100202-12 

"    False  River  near 

New  Roads 

n 

June 

17,  1935 

100420-1 

"    Cane  River  near 

Natchitoches 

it 

June, 

1935 

100529-30  &  9 

Kans.,  Winfield 

a 

Aug. 

12,  1935 

100580 

"       Lake  City 

it 

June, 

1935 

100795  pull. 

Ind.,  Irvington 

O.  P.  Hay 

101386   9  ad. 

Va.,  Seven  Mile  Ford 

A.  Wetmore 

June 

1,  1935 

102705  juv. 

Miss.,  Belzoni 

S.  F.  Hildebrand 

July 

10,  1936 

102911  c?  juv. 

Tenn.,  Reelfoot  Lake 

W.  M.  Perrygo  & 

C.  Lingebach 

May 

8,  1937 

102912 

n              u          u 

W.  M.  Perrygo  & 
C.  Lingebach 

it 

103477-9  juv. 

Vt.,  Swanton 

L.  H.  Babbitt 

June 

21,  1937 

56 


bulletin:  museum  of  comparative  zoology 


107786 

107787 

109178  pull. 
113228  juv. 
115980  pull. 


Term.,  Iron  Creek  near 

Willow  Grove  G.  Gentry 

mouth  of  Wolf 

River 
La.,  Jonesville  G.  K.  Payne 


July    17,  1939 

July    19,  1939 
June,  1940 


Miss.,  Deer  Creek  C.  Hollingsworth        June  24,  1940 


"Amyda  spinifera  aspera1  (Agassiz) 
Plates  17,  18,  19c 

1854.— Trionyx  ferox  WAILES,  Rep.  Agric.  Geol.  Mississippi,  pp.  327,  331 

(Mississippi)  (not  of  Schneider). — Plalypeltis  ferox  AGASSIZ,  Contr. 

Nat.  Hist.  United  States,  2,  1857,  pi.  6,  fig.  3  (Mobile,  Ala.;  Mus. 

Comp.    Zool.,    no.    1608A).— Aspidonectes  ferox    JORDAN,    Man. 

Vert.  North  Amer.,  ed.  5,  1888,  p.  206  (part). 
1857. — Aspidonectes  asper  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  1,  p. 

405   (type  locality,  Lake  Concordia,   La.;  cotypes,  U.S.N.M.  no. 

012349,  Prof.  B.  L.  C.  Wailes,  collector,  and  Mus.  Comp.  Zool,  no. 

37173).— COPE,  Bull.  U.  S.  Nat.  Mus.,  no.  1,  1875,  p.  51  (Lower 

Mississippi  tributaries). — DAVIS  and  RICE,   Illinois  State  Lab. 

Nat.  Hist.,  Bull.  no.  5,  1883,  p.  52.— TRUE,  Fish.  Industr.  United 

States,  sect.  1,  1885,  p.  152.— BAUR,  Amer.  Natural.,  22,  1888,  p. 

1122;  Proc.  Amer.  Philos.  Soc,  31,  July  1893,  p.  217.— BEYER, 

Proc.  Louisiana  Soc.  Nat.  Hist.,  1897-1899  (1900),  p.  43  (Louisiana). 
1892. — Trionyx  agassizii  HAY,  Indiana  Geol.  17  Rep.,  p.  552  (part);  Batr. 

Rept.  Indiana,  1893,  p.  144  (part)  (not  of  Bauer  1888). 
1899. — Aspidonectes  agassizi  JORDAN,  Man.  Vert.  North  Amer.,  ed.  8,  p.  206 

(part  only)  (emendation)  (not  of  Bauer). 

1919. — Amyda  spinifera  BABCOCK,  Mem.  Boston  Soc.  Nat.  Hist.,  8,  no.  3, 
pi.  32  (not  of  text;  not  of  Lesueur).— HALTOM,  Alabama  Mus.  Nat. 
Hist.,  Mus.  Pap.  no.  11,  1931,  p.  142  (Alabama:  Marengo  Co.:  near 
Demopolis,  Tombigbee  River). 

1923.— Trionyx  spiniferus  agassizii  SIEBENROCK,  Verh.  Zool.  Bot.  Ges. 
Wien,  72,  Aug.  1923,  p.  188  (part:  West  Louisiana). 

1939.—  Amyda  spinifera  aspera  STEJNEGER  and  BARBOUR,  Checklist  N. 
Amer.  Amph.  Rept.,  ed.  4,  p.  172  (Lower  Mississippi  tributaries  in 
Louisiana  and  Mississippi) ;  ed.  5,  1943,  p.  213  (Lower  Mississippi  tri- 
butaries in  Louisiana;  rivers  of  Mississippi  and  Alabama). 


1  Latin:  rough;  probably  with  reference  to  the  "prominent  warts  of  the  bony  plates,"  which, 
according  to  Agassiz,  "exist  in  no  other  species  with  which  I  am  acquainted." 


stejneger:  American  soft-shell  turtles  57 


• 


Types  and  type  locality.  The  two  specimens  specifically  mentioned 
by  Agassiz  as  basis  for  his  A.  as  per  were  "an  imperfect  skeleton  .  .  . 
belonging  to  the  Smithsonian  Institution  and  prepared  from  a  speci- 
men forwarded  by  Professor  B.  L.  C.  Wailes  of  Washington,  Missis- 
sippi," and  "a  stuffed  specimen  belonging  to  the  Museum  of  the 
University  of  Oxford,  that  has  been  collected  during  the  Geological 
Survey  of  Mississippi,  under  the  superintendence  of  Professor 
Wailes." 

In  a  letter  dated  Washington,  Miss.,  8:  January,  1853,  Professor 
B.  L.  C.  Waile  wrote  to  Professor  S.  F.  Baird  that  he  had  forwarded 
to  him  a  number  of  reptile  specimens,  among  them  "2  shells  and 
crania  of  Trionyx  ferox."  They  were  entered  in  the  register  of  the 
osteological  collections  of  the  Smithsonian  Institution  under  the 
generic  name  Trionyx  only  by  Professor  Baird  on  March  21,  1853  as 
numbers  1084  and  1086,  received  from'  'B.  L.  C.  Wailes,  Washington, 
Miss."  Of  these  specimens  there  are  now  in  the  National  Museum:  1) 
a  skull  marked  1084;  2)  a  skull,  with  carapace  and  plastron  in  pieces 
(entoplastron  and  epiplastra  missing),  marked  108G;  and  3)  a  carapace 
numbered  12349.  The  specimens  have  no  original  labels  attached  to 
them,  but  the  numbers  on  the  skulls  are  written  on  them  in  ink,  and 
on  the  inside  of  the  carapace  of  No.  12349  there  is  written  with  black 
ink  in  the  same  "professional"  hand  characteristic  of  all  the  specimens 
received  from  Wailes:  "Trionyx  ferox?  Lake  Concordia,  Louisiana, 
1851,  B.  L.  C.  Wailes."  This  is  undoubtedly  the  "imperfect  skeleton" 
examined  by  Agassiz;  it  is  characterized  by  the  "prominent  warts  of 
the  bony  plates"  on  the  posterior  part  of  the  bony  disk  described  by 
Agassiz  (I.e.  p.  406),  which  "bony  warts  exist  in  no  other  species  with 
which  I  am  acquainted."1 

The  second  cotype,  the  "stuffed  specimen"  received  from  the  Uni- 
versity of  Oxford,  Mississippi,  is  in  the  Museum  of  Comparative  Zoo- 
logy (M.C.Z.  no.  37173)  where  I  was  permitted  to  examine  it.  It  is  an 
adult  male,  inscribed  in  the  same  bold,  handsome  style  as  Wailes' 
other  specimens  Trionyx  ferox,  but  no  locality  data.  I  made  the  fol- 
lowing notes:  "Bony  disk  200  mm.  long,  185  mm.  wide;  7  neurals, 
seventh  small;  sculpture  fine,  of  the  spinifer  style,  with  the  bony 
tubercles  on  the  seventh  pleural  described  by  Agassiz,  tubercles  on  the 
posterior  and  anterior  flaps  of  leathery  disk,  spinous  tubercles  on  edge 
of  carapace  anteriorly  between  legs;  callosities  on  plastron  large,  al- 
most meeting  on  the  mid-line;  a  median  large  triangular  callosity 

1  Agassiz  did  not  know  the  bony  disk  of  the  true  A.  ferox  from  Florida. 


58  bulletin:  museum  of  comparative  zoology 

(sides  about  25  mm.)  on  the  entoplastron ;  entoplastral  angle  slightly 
less  than  90°. 

The  specimens  which  Agassi z  mentions  as  having  been  received 
"through  the  kindness  of  Mr.  Winthrop  Sargent  of  Natchez"1  may  be 
regarded  as  paratypes.  One  of  them,  a  very  large  male  (M.C.Z.  no. 
1597),  I  have  examined  and  made  the  following  measurements  and 
notes : 

Total  length  of  leathery  carapace  450  mm. 

Width  of  leathery  carapace  370 

Length  of  bony  carapace  240 

Width  of  bony  carapace  (at  pleural  4)         240 
Height  of  body  (at  neural  1)  88 

Large  rounded  tubercles  on  front  edge  of  leathery  carapace;  flat 
tubercles  of  about  same  size  on  front  and  hind  flaps;  callosities  on 
xiphi-plastra  meeting;  closest  approach  of  hyo-hypoplastral  callosities 
8  mm. ;  bony  ridges  comparable  to  those  on  bony  carapace  of  U.S.N.M. 
no.  1086;  sculpture  somewhat  coarser  (because  of  larger  size  of  speci- 
men) ;  entoplastral  angle  about  90°.  Distinct  traces  of  two  black  mar- 
ginal rings  on  hind  flap  of  carapace. 

A  young  paratype,  probably  of  the  same  origin  (M.C.Z.  no.  1622) 
with  a  recent  label  in  lead  pencil  "No.  1622.  Type  Amyda  asper 
(Agassiz)  Lake  St.  John,  Miss.2  W.  Sargent  leg.  et  don."  measures  ap- 
proximately 65  mm.  in  length  and  57  mm.  in  width.  The  secondary 
marginal  lines  on  the  posterior  flap  of  the  carapace  characteristic  of 
normal  individuals  of  the  subspecies  of  corresponding  age  are  repre- 
sented by  two  series  of  closely  set  spots  (pi.  16,  fig  1). 

Notes  on  synonymy.  Special  attention  is  called  to  the  reference  to 
Agassiz's  illustration  (Contrib.,  2,  pi.  6,  fig.  3)  of  a  young  specimen  of 
A.  aspera  under  the  name  PlatypeHis  fcrox,  as  it  apparently  has  given 
rise  to  great  confusion  among  southern  herpetologists  who,  because  of 
it,  have  mistakenly  identified  young  specimens  from  Alabama,  Missis- 
sippi, and  Louisiana.  The  young  of  the  true  A.  ferox  was  unknown  to 
Agassiz,  but  on  account  of  the  locality  (Mobile)  of  the  specimen  figured 
and  its  similarity  in  color  pattern  to  certain  soft-shell  turtles  from 
South  Carolina  he  assumed  that  it  represented  the  Florida  species. 
The  original  of  the  figure  3  appears  to  be  still  at  the  Museum  of  Com- 
parative Zoology  bearing  the  number  1608A.    It  turns  out  to  be  a 

'Agassiz,  in  1856,  had  J.  Burkhart  make  colored  illustrations  from  a  female  of  the  same  lot 
of  specimens  which  shows  plainly  the  marginal  black  rings  on  the  posterior  flap.  Reduced 
copies  of  the  drawings  are  herewith  presented  (pis.  20,  21)  through  the  kind  permission  of 
Dr.  Thomas  Barbour,  Director  of  the  Museum  of  Comparative  Zoology. 

2  Possibly  a  slip  for  Louisiana. 


stejneger:  American  soft -shell  turtles  59 

young  of  Agassiz's  own  Aspidonectes  as  per,  and  the  figure  is  herewith 
reproduced  under  its  proper  name  (pi.  19,  fig.  c)  for  comparison  with 
the  young  of  the  true  A.  ferox  (pi.  19,  fig.  a  and  b). 

Babcock's  figure  of  A.  spinifera,  as  quoted  above,  does  not  represent 
the  true  nominate  form  of  that  species,  but  is  either  an  A.  spinifera 
aspera  or  an  A.  agassizii.  Dr.  Babcock,  in  a  letter  of  December  13, 
1933,  kindly  informs  me  that  the  picture  was  made,  during  his  absence 
in  1918,  from  a  living  specimen,  the  origin  and  disposition  of  which 
he  has  been  unable  to  trace.  The  exact  identification  depends  some- 
what on  the  artist's  accuracy.  My  best  guess  is  that  the  specimen  was 
an  A.  aspera,  judging  by  the  extreme  length  of  the  proboscis. 

Variation.  A  series  of  four  living  young  adults  were  presented  in 
September,  1934,  by  Mr.  S.  E.  Brand,  of  Canton,  Mississippi,  all  taken 
in  barrow  pits,  three  on  Pearl  River  and  one  on  Big  Black  River.  The 
specimens  unquestionably  represent  the  same  form  and,  as  highly 
instructive,  some  of  the  notes  made  at  the  time  may  be  of  interest,  the 
color  designations  in  quotation  marks  having  reference  to  Ridgway's 
Nomenclature  of  Colors,  1886: — 

U.S.N.M.  no.  95191,  9  ad.  Iris  clear  "primrose  yellow"  with  black 
horizontal  bar  not  quite  reaching  pupil.  General  color  above  "tawny- 
olive,"  head,  neck,  and  legs  densely  speckled  with  dark  brown,  cara- 
pace with  obscure,  irregular  blotches  of  "raw  umber,"  in  the  center 
of  which  one  or  more  small  blackish  spots  form  interrupted  ocelli,  the 
two  outer  rows  of  which  posteriorly  assume  the  form  of  short  lines 
parallel  with  the  submarginal  ring;  marginal  edge  dull  "olive-buff," 
top  of  head  like  carapace  with  the  shallow  fork  on  the  snout  chiefly 
indicated  by  the  black  outlines;  side  of  head  like  top  down  to  the  lips; 
a  slightly  paler  postocular  band  tinged  with  yellow  and  narrowly  but 
very  distinctly  margined  with  black,  the  upper  line  continuing  back- 
ward the  whole  length  of  the  neck ;  upper  side  of  legs  slightly  paler  than 
carapace  and  more  tinged  with  olive;  under  side  pale  flesh  color  caused 
by  the  fine  network  of  blood  vessels  shining  through  the  white  ground 
color;  plastral  callosities  pale  "vinaceous-cinnamon"  more  or  less 
tinged  with  bluish;  under  side  of  feet  tinged  with  "verditer  blue"  with 
a  wash  of  yellow  deepening  towards  the  outer  margin,  inner  half 
spotless,  outer  half,  including  web,  marked  with  heavy  blackish  anas- 
tomozing  lines  and  spots;  claws  yellowish  white;  under  side  of  neck 
faintly  mottled  with  obscure  "pinkish  vinaceous"  spots;  throat 
strongly  washed  with  "pale  blue";  lips  and  under  side  of  proboscis 
tinged  with  "gallstone-yellow."  Tubercles  on  anterior  edge  of  cara- 
pace triangular,  pointed,  about  3  mm.  long;  anterior  flap  densely 


GO  bulletin:  museum  of  comparative  zoology 

crowded  with  smaller  and  blunter  tubercles  of  various  sizes;  posterior 
flap  with  larger,  oblong,  blunt  tubercles,  fairly  regularly  spaced;  cara- 
pace posteriorly  with  similar  but  lower  and  longer  tubercles  in  fairly 
regular  longitudinal  series.  Skin  covering  nuchal  of  bony  carapace 
with  a  median  longitudinal  series  of  about  4  enlarged  tubercles  ex- 
tending on  to  anterior  leathery  flap.  Fontanelles  between  nuchal  and 
first  pair  of  pleurals  covered  with  smooth  skin  upon  which  are  a  few 
tubercles.  Callosities  covering  plastral  bones  rather  small,  leaving 
median  fontanelles  larger  than  xiphiplastral  callosities;  no  trace  of 
callosities  on  entoplastron  and  ectoplastron ;  entoplastral  angle  90°. 

U.S.N.M.  no.  95192,  also  from  a  "barrow  pit"  at  the  nearby  Big 
Black  River,  a  young  adult  9  ,  smaller  than  95191,  is  structurally  al- 
most identical  with  it,  except  for  the  smaller  dimension  and  being 
slightly  broader.  The  coloration  of  the  living  specimen  was  darker  and 
more  pronounced.  Iris  "ochraceous  buff,"  black  horizontal  bar  reach- 
ing pupil.  Carapace  "raw  umber"  with  large  irregularly  buffy-edged 
blotches  of  "mummy  brown,"  increasing  towards  and  infringing  upon 
the  marginal  "buffy"  edge  and  its  bordering  dusky  ring;  towards  the 
margin  the  blotches  coalesce  more  or  less  so  as  to  form  three  distinct 
but  interrupted  submarginal  rings;  color  of  upper  side  of  neck,  head, 
and «legs  like  that  of  the  carapace  but  more  "tawny"  and  thickly 
sprinkled  with  heavy  blackish  spots ;  forked  figure  on  top  of  snout  deep 
"ochraceous  buff,"  with  very  distinct  black  edges  like  the  ochraceous 
postocular  stripe;  under  side  milk  white  strongly  suffused  with  flesh- 
color;  palms  and  soles  strongly  tinged  with  tawny,  palms  heavily 
marked  with  coarse  blackish  anastomozing  lines  and  spots,  soles  al- 
most devoid  of  markings,  claws  white,  throat  tinged  with  pale  blue; 
callosities  "vinaceous-cinnamon"  with  slight  tinge  of  blue  in  center. 

U.S.N.M.  no.  95193,  young  adult  d\  from  Pearl  River,  differs 
structurally  from  the  two  described  females  in  showing  no  trace  of 
fontanelles  on  carapace,  in  much  larger  plastral  callosities  than  on 
xiphiplastra  nearly  meeting  in  the  middle,  those  covering  hyoplastra 
and  hypoplastra  only  4  mm.  apart;  in  addition  there  is  a  large  triangu- 
lar callosity  on  the  entoplastron ;  the  row  of  tubercles  on  the  edge  of 
anterior  flap  of  carapace  barely  indicated  and  no  tubercles  on  flap  and 
nuchal.  In  addition,  the  skin  of  the  carapace,  bony  disk  as  well  as 
lateral  and  posterior  flaps,  is  densely  sprinkled  with  very  minute  hard 
tubercles  which  make  the  skin  feel  like  fine  sandpaper.  The  color 
notes  on  the  living  specimens  are  as  follows :  Iris  pale  buff,  black  bar 
scarcely  separated  from  pupil.  Upper  side  nearly  uniform  "tawny 
olive,"  the  region  of  the  bony  disk  distinctly  more  olive;  small,  round, 


stejneger:  American  soft-shell  turtles  61 

dark  brown  spots  of  varying  size  scattered  sparingly  over  the  carapace; 
the  normal  margin  not  perceptibly  lighter  than  rest  of  carapace  and 
barely  set  off  from  it  by  the  obscure  dusky  submarginal  ring;  no  in- 
dications of  additional  rings  by  dark  lines  or  spots;  top  of  head  and 
neck  like  back  with  minute  black  scattered  dots;  lips  and  sides  of  neck 
strongly  washed  with  "gallstone  yellow,"  with  normal  postocular 
stripe  pattern  indicated;  fork  figure  on  snout  normal,  buff  colored, 
black-edged;  legs  above  and  feet  coarsely  spotted  with  black;  upper 
side  of  tail  like  feet  with  lateral  blackish  lines  converging  backwards. 

U.S.N.M.  no.  95194,  from  the  same  locality,  is  also  a  male,  and 
slightly  smaller.  It  shares  the  same  structural  characteristics,  only 
the  tubercles  on  the  anterior  edge  of  carapace  are  slightly  larger  and 
more  distinct;  the  outline  of  the  soft  carapace  is  more  oval  than 
rounded  ovate;  the  xiphiplastral  callosities  are  continuous,  but  the 
distance  between  the  hyohypoplastral  callosities  is  a  trifle  wider,  and 
the  entoplastral  callosity  somewhat  smaller;  the  "sandpaper"  effect 
of  the  back  is  very  much  alike,  but  a  double  series  of  larger  flat  tuber- 
cles on  the  mid-line  of  the  7th  pleural  is  quite  noticeable.  The  colora- 
tion is  also  much  the  same,  only  the  ground  color  of  the  carapace  is 
lighter,  and  the  dark  spots  distinct,  the  chief  difference  being  the  inter- 
rupted rows  of  the  three  marginal  rings;  on  the  other  hand,  the  dusky 
pattern  elsewhere  is  finer;  the  pale  edge  of  the  leathery  disk  is  slightly 
paler  than  the  rest  of  the  disk. 

For  an  easier  appreciation  of  the  more  striking  variations,  they  are 
summarized  in  table  7.  The  differences  are  of  various  significance.  As 
certainly  indication  of  sex  is  the  greater  development  of  the  plastral 
callosities  of  the  males,  besides  the  well-known  difference  in  the 
length  of  the  tail,  the  "sandpaper"  effect  seems  to  be  peculiar  to  a  cer- 
tain age  (or  size,  or  season?)  of  young  males;  the  greater  development 
of  the  tubercles  or  "spines"  may  be  correlated  with  sex  or  with  age  or 
with  both,  remembering  that  the  females  are  larger  than  the  males; 
the  closing  of  the  nuchal  fontanelles  of  the  carapace  seems  to  take 
place  much  earlier  in  the  males  than  in  the  females;  the  slight  differ- 
ence in  the  outline  of  the  leathery  disk  (as  seen  from  above)  is  not  due 
to  sex,  and,  at  the  stage  of  the  specimen  under  discussion,  not  to  age, 
though  the  general  rule  in  these  turtles  is  that  the  young  are  more 
circular  than  the  old  ones.  None  of  the  above  characters  seem  to  be 
of  specific  or  subspecific  significance.  On  the  other  hand,  the  presence 
of  two  or  more  concentric  blackish  rings  inside  the  normal  Amydan 
dusky  ring  delimiting  the  pale  rim  of  the  leathery  carapace  is  of  diag- 
nostic value  in  defining  the  subspecies,  but  unfortunately  it  has  a 


62 


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tendency  to  be  obscured  or  absorbed  with  age.  Individual  specimens 
may  occasionally  be  found  without  these  additional  rings  among  nor- 
mal populations,  but  they  are  the  "intermediates"  which  justify  the 
use  of  the  trinominal  nomenclature.  On  the  other  hand,  it  is  significant 
that  among  a  very  large  number  of  specimens  from  northernmost 
Montana  to  Louisiana  and  from  Colorado  to  Lake  Champlain,  I  have 
not  seen  an  authentic  specimen  of  Amy  da  spinifer  with  two  or  more 
submarginal  rings. 


Geographical  distribution 

Lower  Mississippi  tributaries   in   Louisiana  and  Mississippi   and 
Alabama. 


Table  7 
Variations  in  young  adult  A.  aspera  from  Mississippi 


Mississippi 


Pearl 

Black 

Pearl 

Pearl 

River 

River 

River 

River 

95191 

95192 

95193 

95194 

Sex 

9 

9 

& 

o* 

Soft  carapace  length                               mm. 

286 

210 

180 

158 

width                                 mm. 

242 

182 

152 

140 

Bony  carapace  length                            mm. 

173 

119 

111 

99 

width                             mm. 

131 

85 

110 

98 

Plastral  callosities 

Small 

Small 

Large 

Large 

Callosity  on  entoplastron 

0 

0 

+ 

+ 

Spines  on  edge  of  carapace 

Large 

Median 

Small 

Small 

pointed 

pointed 

blunt 

blunt 

Tubercles  on  median  line  of  front  carapace 

flap 

+ 

+ 

0 

0 

Carapace  skin  "sandpapered" 

0 

0 

+ 

+ 

Carapace  fontanelles 

+ 

+ 

0 

0 

Outline  of  carapace  (from  above)  oval 

+ 

+ 

"     "           "             "         "       ovate 

+ 

+ 

stejneger:  American  soft-shell  turtles 


63 


Intergrades  aspera-spinifera 

It  has  been  repeatedly  asserted  that  the  only  differences  by  which 
specimens  of  A.  asper  a  can  be  recognized  in  the  adult  state  are  "the 
very  coarse  and  large  tubercles  of  the  front  and  hind  part  of  the  cara- 
pace, which  extend,  behind,  even  over  the  bony  shield,  and  are  there 
supported  by  prominent  warts  of  the  bony  plates"  (Agassiz,  Contr., 
1,  p.  406)  and  by  the  fact  "that  in  younger  specimens  of  Asp.  asper 
there  are  .  .  .  two  or  three  black  lines  separating  the  pale  rim  of  the 
posterior  margin,  whilst  there  is  only  one  in  Asp.  spinifer."  (Agassiz, 
I.e.).  As  Agassiz  himself  observes,  these  lines  fade  away  "pretty  soon." 

Table  8 
Cranial  measurements  of  A.  asper  a  in  millimeters 


Basicranial  length 


mm. 


Tip  of  snout  to  orbit 
Horizontal  diameter  of  orbit 
Orbit  of  tympanic  cavity 
Longest  diameter  of  temporal  fossa 
Interorbital  width 
Width  of  maxillary  alveole 
Length  of  internal  choanae 
Internal  choanae  to  intermaxillary 

foramen 
Length  of  intermaxillary  foramen 
Length  of  mandibular  symphysis 
Width  of  mandibular  alveole  surface 


C 
o 

G 

A 

to 

oj 


CO 
00 

o 


63       63 


16 
12 
19 
11 

5 

5.5 
10 

7 

9 


a 


oo 

o 


16 
10.5 
18.5 
12 

6 

5 

9 

6.5 
6.2 


03 

c 

o 
>- 

0> 

a 

C 
o3 


O 
i— i 

ec 

OS 

o 


63 

16 
12 
19 
13 

5.5 

5 

9 

6 

6.5 

9 

4 


0) 

o 

"oj 

c 


3 

O 

1-5 


o 

00 
CD 


37 


10 
8 
9 

6.5 
3 
3 
6 

6 
5 
6 
2.5 


cj 


Z 


a 


CO 


23       69 


6 

5.5 

6 

4 

2 

2 

5 

3 
3 
3 
2 


o3 
C 
c3 


3 
O 

h-1 


CO 
CO 
<M 
Oi 
CM 

o 


19 
13 
20.5 
13 

5 

6 
10 

6.5 
7 
11.5 
5.5 


m 

C 

s, 


o 

a 

02 
CO 


bD 
cj 
s-, 

> 

< 


53.0 

13.8 
10.1 
15.3 
9.9 
4.4 
4.4 
8.1 

5.9 
5.5 
6.4 
3.5 


64 


bulletin:  museum  of  comparative  zoology 


Specimens  of  intermediate  ages  are  difficult  to  identify  with  our  present 
knowledge.  Apparently  the  double  or  triple  rings  are  not  always  a 
constant  character.  I  have  before  me,  through  the  courtesy  of  Miss 
Fannye  A.  Cook,  a  specimen  collected  about  a  mile  or  two  southeast 
of  Brookhaven,  Lincoln  Co.,  Mississippi,  in  a  small  tributary  of  the 
Bogue  Chitto,  Pearl  River  drainage.  Its  carapace  measures  only  43 
mm.  in  length,  hence  it  is  quite  young.  It  seems  to  be  an  aberrant 
specimen  of  asper  a,  the  anomaly  being  in  the  absence  of  the  second 
dark  ring  on  the  carapace  margin.  From  its  locality  it  ought  to  be 
aspera,  but  such  abnormal  (or  "incompleted"  or  "reversed")  specimens 
are  known. 


List  of  specimens  in  the  U.  S.  National  Museum 


01084 

Miss.,  Washington? 

B.  L.  C.  Wailes 

01086 

H                             11 

ii 

Cotype  of 
Aspidenectes  asper 
Agassiz 

7653-4 

"      Monticello 

H.  Tennison 

012349 

La.,  Lake  Concordia 

B.  L.  C.  Wailes 

1851  Cotype  of 
Aspidonectes  asper 
Agassiz 

13250  9 

"    New  Orleans 

R.  W.  Shufeldt 

1883 

029266 

u 

S.  W.  Harvey 

029310  ad.  9 

"    near  New  Orleans 

U.  S.  Fish  Comm. 

66147  juv. 

"    Madisonville 

? 

May  29,  1886 

68054  cf  adol. 

"    Roberts 

R.  F.  Shaw 

79350-1 juv. 

Miss.,  1  mi.  W.  of  Mel- 

R.  Kellogg  and 

vin 

N.  Boss 

Oct.  1929 

83996  9  juv. 

Ala.,  3  mi.  S.E.  of 

Coatopa 

C.  E.  Burt 

July     1,  1931 

95191  9  ad. 

Miss.,  Pearl  River 

S.  E.  Brand 

Aug.  1934 

95192  9  adol. 

"      Big  Black  River 

ii 

u 

95193  d"  adol. 

"      Pearl  River 

ii 

ii 

95194  d"  adol. 

u               ii 

a 

ii 

100650  c?  ad. 

La.,  near  Atchafalaya 

C.  E.  Burt 

June  17,  1935 

100805  pull. 

Miss.,  Enterprise 

O.  P.  Hay 

1881 

115979  9  adol. 

"      near  Guntoun 

E.  &  W.  H.  Patten 

Aug.    4,  1940 

115981 juv. 

"      Chookatonkchia 

Creek 

H.  L.  Owens 

June  30,  1941 

stejneger:  American  soft-shell  turtles  65 

Amyda  emoryi1  (Agassiz) 
Plates  24-25 

1849.     Trionyx  ferox  ROEMER,  Texas,  p.  171,  p.  459  (at  New  Braunfels, 
Texas,  in  the  Guadalupe  and  Comal  Rivers). 

1857. — Aspidonectes  emoryi  AGASSIZ,  Contr.  Nat.  Hist.  United  States  1,  p. 
407;  2,  pi.  6,  figs.  4-5  (type-locality,  Rio  Grande  River,  near  Browns- 
ville, Texas;  cotypes,  U.  S.  Nat.  Mus.  No.  7855;  Mus.  Comp.  Zool., 
Nos.  1909,  1913;  Dr.  Kennedy,  collector;  Williamson  Co.,  Texas). — 
COPE,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  1875,  p.  51  (Texas);  No.  17, 
1880,  p.  13  (Dallas;  Helotes  Creek,  near  San  Antonio,  Texas). — 
TRUE,  Bull.  U.  S.  Nat.  Mus.,  No.  24,  1883,  p.  29  (Matamoras, 
Mexico;  Texas:  Brownsville,  Rio  Grande,  Rio  Seco,  Braunfels;  Old 
Fort  Cobb,  Oklahoma);  Fish.  Industr.  United  States,  sect.  1,  1884, 
p.    152.— BEYER,   Proc.    Louisiana  Soc.    Nat.,    1897-1898,   p.   43 
(Louisiana).— STRECKER,  Trans.  Texas  Acad.  Sci.,  4,  pt.  2,  no. 
5,  p.  6  (McLennan  Co.,  Texas);  Proc.  Biol.  Soc.  Washington,  21, 
March  21,  1908,  p.  79  (Bazos,  Bosque  River,  McLennan  Co.,  Texas, 
abundant);  23,  1910,  p.  121  (Bullhide  Creek,  McLennan  Co.,  Texas). 
Trionyx  emoryi  BOULENGER,  Cat.  Chel.  Brit.  Mus.,  1889,  p.  258  — 
STRAUCH,  Mem.  Acad.  Sci.  St.  Petersbourg,  ser.  7,  38,  no.  2,  1890, 
p.  117  (Texas).— DITMARS,  Reptile  Book,  1907,  p.  78  (tributaries 
of  the  Rio  Grande  in  Texas  and  Mexico).— SIEBENROCK,  Zool. 
Jahrb.,  Suppl.  10,  pt.  3,  1909,  p.  603  (Colmisneil,  Tyler  Co.,  South 
Bosque  Riv.,  Texas);  Verh.  Zool.  Bot.  Ges.  Wien,  73,  1923,  p.  190.— 
LINSDALE  and  GRESSITT,  Copeia,  1937,  no.  4,  Dec.  31,  pp.  222- 
225,  figs.  1-3  (Colorado  River:  Delta,  Lower  California;  Clark  Co., 
Nevada;   Mohave  Co.,   Arizona;  California  Lakes,   Imperial   Co., 
California;  transpl.  to  Colorado  Riv.?). — LINSDALE,  Proc.  Ameri- 
can Acad.  Arts  Sci.,  73,  no.  8,  May  1940,  p.  255  (Nevada:  Clark  Co. : 
Colorado  River). 
Amyda  emoryi  STEJNEGER  and  BARBOUR,  Checklist  North  Amer. 
Amph.  Rept.,  ed.  1,  1917,  p.  124  (Rivers  of  Texas,  north  into  south- 
ern Oklahoma  and  Arkansas);  ed.  2,  1923,  p.  140;  ed.  3,  1933,  p.  153. 
-PRATT,  Man.  Vert.  United  States,  1923,  p.  249.—  ?  HAY,  Pan- 
Amer.  Geol.,  39,  March  1923,  p.  119,  pi.  9,  figs.  2-4  (fossil,  Brazos 
River  at  Pittbridge,  Texas).— SCHMIDT,  Copeia,  no.  131,  June  30, 
1924,   p.   64    (Arizona;  introduced).— STRECKER,   Baylor  Univ. 
Bull.,  27,  no.  3,  Sept.  1924,  p.  47  (eastern  Oklahoma);  Contr.  Baylor 
Univ.  Mus.,  no.  2,  Jan.  15,  1926,  p.  3  (Somervell  Co.,  Texas);  no.  3, 
Feb.  15,  1926,  p.  4  (Liberty  Co.,  Texas);  no.  6,  June  15,  1926,  p.  8 

1  To  Col.  Wm.  H.  Emory,  U.S.A.,  under  whose  command  part  of  the  type  material  was  col- 
lected, "I  take  great  pleasure,  therefore,  in  dedicating  this  species  to  that  distinguished  officer." 

(Agassiz) 


66  bulletin:  museum  of  comparative  zoology 

(Cibolo  Creek,  Boerne,  Tex.);  no.  7,  July  15,  1926,  p.  7  (Cedar  Creek, 
Henderson  Co.,  Tex.);  no.  19,  1929,  p.  15  (Trinity  Riv.,  Ft.  Worth, 
Tex.);  no.  23,  June,  1931,  p.  16  (Colorado  Riv.,  Trevis  Co.,  Tex.); 
Baylor  Bull.,  38,  no.  3,  Aug.  1935,  p.  23  (Texas:  Cibolo  Creek),  p.  32 
(Texas:  Real  Co.).— ORTENBURGER,  Proc.  Oklahoma  Acad.  Sci., 
6,  pt.  1,  1926,  p.  100  (LeFlore  Co.,  Oklahoma).— STRECKER  and 
WILLIAMS,  Contr.  Baylor  Univ.  Mus.,  no.  12,  Dec.  27,  1927,  pp. 

11,  15  (San  Marcos  and  Blanco  Rivers,  Texas).— RUST,  Blatt. 
Aquar.  Terrarienk.,  45,  1934,  p.—,  sep.,  p.  12.— LITTLE  and  KEL- 
LER, Copeia,  1937,  no.  4,  Dec.  31,  pp.  216,  221  (Mesilla  Valley, 
Dona  Ana  Co.,  New  Mexico). — GAIGE,  Univ.  Michigan  Stud.  Sci., 

12,  1937,  p.  304  (Mexico,  Tamaulipas,  Rio  Purificaci6n,  N.  of  Ciudad 
Victoria).— DELLINGER  and  BLACK,  Occas.  Pap.  Univ.  Arkansas 
Mus.,  no.  1,  June  1938,  p.  46  (?  Arkansas:  Salina  Riv.  near  Benton? 
[probably  Texas,  U.S.N.M.  no.  17823-L.S.]).— SMITH,  Ann.  Car- 
negie Mus.  Pittsburgh,  27,  1939,  p.  312  (Mexico:  Tamaulipas; 
Nuevo  Laredo). 

1870. — Aspidonectes  emyda  "Agassiz,"  GRAY,  Suppl.  Cat.  Shield  Rept.  Brit. 

Mus.,  pt.  1,  p.  95  (lapsus). 
1870. — Aspidonectes  georgii  "Agassiz,"  GRAY,  Suppl.  Cat.  Shield  Rept.  Brit. 

Mus.,  pt.  1,  p.  109  (lapsus). 

1893.  Platypeltis  emoryii  BAUER,  Proc.  Amer.  Philos.  Soc,  31,  p.  220  (emen- 
dation).— Amyda  emoryii  STRECKER,  Copeia,  no.  162,  1927,  p.  9 
(food  habits);  Contr.  Baylor  Univ.  Mus.,  no.  15,  July  10,  1928,  p.  6 
(Bosque  Riv.  near  Valley  Mills,  Bosque  Co.,  Texas);  no.  16,  Aug.  4, 
1928,  p.  21  (Texas:  vernacular  names). 

Agassiz  (p.  407),  as  character  aiding  in  identifying  this  species, 
calls  attention  to  skin  of  the  carapace  being  "dotted  all  over  with 
small  whitish  tubercles  like  grains  of  sand."  This  is  not  a  specific 
character,  it  seems  to  be  a  condition  of  the  skin  due  to  season  or  age,  as 
similarly  "sandpaper"  specimens  are  encountered  in  several  of  the 
species. 

Skull.  The  skulls  of  A.  emoryi  and  A.  svinifera  are  very  much 
alike.  The  snout  in  emoryi  is  slightly  shorter  and  somewhat  broader 
anteriorly,  the  nasal  cavity  relatively  shorter  and  the  angle  formed  by 
the  anterior  processes  of  the  prefrontal  bones  more  obtuse.  The 
alveolar  surface  of  the  maxillaries  are  somewhat  wider.  In  these  re- 
spects the  emoryi  are  even  closer  to  A.  fero.v. 

As  the  differences  in  the  skull  between  A.  spinifera  and  emoryi  are 
very  slight,  many  skulls  can  hardly  be  told  apart.  In  the  former  the 
choanae  and  the  intermaxillary  foramen  average  a  trifle  larger.  The 
orbit  in  A.  spinifera  is  also  placed  slightly  more  backward  on  the 


STEJNEGER:    AMERICAN    SOFT-SHELL   TURTLES  67 

average  than  in  A.  emoryi.  As  a  consequence  the  snout  appears  a 
mere  trifle  longer.  However,  it  is  difficult  to  understand  how  Boulen- 
ger  (Cat.  Chel.  Brit.  Mus.,  18S9,  pp.  245-246)  came  to  diagnose  A. 
emoryi  as  having  "the  snout  (on  the  skull)  obtuse,  hardly  as  long  as  the 
diameter  of  the  orbit,"  and  the  other  two  {spinifera  and/<  rox)  having 
it  "a  little  longer."  In  26  skulls  measured  by  me  the  horizontal  diame- 
ter of  the  orbits  in  ferox,  spinifera,  and  emoryi  averages  20.0,  19.6,  and 
19.1  mm  respectively  and  the  snout  (as  measured  from  the  orbit) 
24.9,  26.5,  and  24.4  mm.  In  the  six  emoryi  measured  by  me  the  snout 
is  2.5,  3.0,  4.0,  5.5,  1.5,  and  2.0  mm  longer  than  the  ort.it.  The  relative 
dimensions  of  snout  and  orbit  are  therefore  unavailable  as  a  diagnostic 
character.  It  should  be  noted,  finally,  that  in  emoryi  the  alveolar 
surface  of  the  maxilla  of  medium  sized  skulls  is  slightly  wider  than  in 
the  other  two,  but  too  slightly  and  variably  so  to  be  of  much  help  in 
diagnosing. 

Plastron.  Referring  to  Siebenrock's  remarks  about  the  plastron  of 
.4.  emoryi  (Sitz.  Ber.  Akad.  Wiss.  Wien,  Math.  Nat.  KL,  111,  1902,  p. 
830)  it  is  well  to  note  that  the  bones  are  essentially  as  in  A.  spinifera. 
The  entoplastral  angle  in  adults  seems  to  be  more  acute.  The  callosi- 
ties appear  to  be  more  developed  in  a  male  (U.S.N.M.  no.  26426); 
they  are  almost  as  large  as  in  the  A.  mutica  figured  by  him  (p.  823,  fig. 
5),  but  of  course  without  trace  of  callosities  on  entoplastron  or  epi- 
plastra.   The  anterior  portion  of  the  epiplastra  is  rather  long. 

Color  of  live  specimens 

Two  adult  males  from  Houston,  Texas,  are  colored  as  follows.  Xo. 
94335  has  the  top  of  head  and  dorsal  aspect  of  neck  dark  olive  green, 
becoming  gradually  more  green  on  posterior  half  of  neck.  Carapace, 
bony  disk  Van  Dyke  brown,  bistre  on  the  soft  parts,  with  irregular 
blackish  brown  anastomozing  spots.  Sides  of  neck  almost  citron  yellow 
fading  into  whitish  on  the  middle  line  of  the  ventral  aspect  of  the  neck 
which  anteriorly  changes  into  verditer  blue  on  the  throat,  darkening 
to  almost  indigo  blue  on  mentum  and  outer  half  of  lower  jaw  and 
tympanic  region,  and  extending  a  little  below  and  backwards  on  side 
of  neck;  the  yellow  of  the  side  of  the  neck  fades  into  a  dull  olive  yel- 
low with  a  few  scattered,  almost  blackish  spots. 

Tubercles  on  anterior  edge  of  soft  carapace  large  (3-4  mm.),  tri- 
angular, spaced  apart  by  the  width  of  their  bases. 

Underside  white,  also  feet,  but  fingers  and  webs  pale  (dull)  sage 
green  with  obscure  dark  marblings. 


68  bulletin:  museum  of  comparative  zoology 

Their  measurements  differ  as  follows: 

d*  ad.  94335  &  ad.  94336 

Soft  carapace,  long                         342  mm.  354  mm. 

286    "  288    " 

Height  of  body                                 86     "  84     " 

Tip  of  tail  beyond  carapace            40  32 

In  No.  94336  the  colors  are  essentially  as  in  the  other  except  that 
on  head  and  neck  the  yellow  is  a  little  deeper  and  that  there  are  a 
large  number  of  small  blackish  spots  on  -sides  of  head,  even  including 
lower  eyelid,  lips,  and  base  of  proboscis,  and  definite  blackish  lines 
running  from  eye  obliquely  backward  to  base  of  lower  jaw  which 
is  also  outlined  by  similar  lines;  dusky  obscure  large  (average  15 
mm.)  ocellar  markings  on  under  side  of  neck.   Tubercles  like  94335. 

Two  specimens  collected  by  Dr.  and  Mrs.  A.  H.  Wright,  U.  S.  N.  M. 
nos.  94456-7  are  colored  as  follows.  In  no.  94456  the  neck  and  iris 
chrome  yellow  more  or  less  pale,  iris  with  horizontal  black  lens. 
Carapace  above  uniform  wood  brown.  Marginal  edge  washed  with 
chromium  green.  Upper  soft  parts  of  head  and  legs  pale  chromium 
green  washed  with  wood  brown  on  top  and  sides  of  head.  Tail  above 
white,  with  posterior  central  part  dull  chromium  green  washed  with 
cadmium.  Tubercles  on  carapace  pale  buff,  minute,  round,  densely 
scattered  over  the  whole  carapace,  hand,  and  soft  parts.  Underside 
white.  Neck  underneath  very  pale  cadmium  shading  anteriorly  into 
pale  verditer  blue  washed  with  pale  cadmium  (on  throat).  Dark 
markings  on  throat  strongly  tinged  with  bluish.  Fleshy  lips  pale 
cadmium  yellow.  Pale  cadmium  yellow  on  fork  of  snout  fading  away 
anteriorly.    Triangle,  base  between  eyes  slightly  angular. 

No.  94457  has  the  same  data,  but  a  dark  ring  of  minute  blackish 
specks  surrounding  tubercles.  Underside  of  foot  same  color  as  upper 
chromium  green  parts — spotted. 


stejneger:  American  soft-shell  turtles 


69 


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70  bulletin:  museum  of  comparative  zoology 

Another  male,  U.S.N.M.  104240,  from  Pecos  River,  near  Dryden, 
Terrell  Co.,  Texas.   F.  M.  Setzler,  coll.  shows  still  further  variation. 

Anterior  border  of  disk  very  obscurely  tuberculated.  Skin  on  cara- 
pace shagreened  like  bony  disk.  Skin  on  flaps  smooth,  leathery:  a 
round  smooth  area  over  each.  The  fontanelle  smooth.  Part  of  flap 
behind  bony  disk  with  a  regular  pattern  of  whitish  "pimples"  or 
tubercles,  those  along  the  middorsal  line  in  pairs.  No  "sandpaper" 
effect.  Palms  and  soles  yellowish  white,  unspotted.  Carapace  (day 
or  two  after  death) :  flap  Isabella  color  (Ridgway,  pi.  Ill,  fig.  23);  bony 
disk  more  tawny  olive  (fig.  17);  neck  and  limbs  above  pale  olive  gray 
like  pickled  unripe  olives,  ventral  half  of  neck  yellowish  white  with  a 
faint  trace  of  a  similarly  colored  band  from  lower  edge  of  eye  back- 
wards along  the  neck.  All  upper  soft  parts  with  numerous  black  dots 
more  or  less  arranged  in  longitudinal  series. 

Geographical  distribution 

Rivers  of  Texas,  north  into  southern  Oklahoma  and  Arkansas,  west 
(introduced?)  to  southeastern  California,  the  adjacent  portion  of 
Lower  California,  and  Clark  County,  Nevada;  east  to  western  Louisi- 
ana; northern  Mexico. 

List  of  specimens  in  the  U.  S.  National  Museum 


7176 

Okla.,  Old  Ft.  Cobb 

E.  Palmer                  May 

4,  1868 

7614-20, 

Mex.,  Matamoras, 

7662-25 

Tamaulipas 

L.  B.  Couch 

7628-33 

Mex.,  Matamoras, 

Tamaulipas 

n 

7635-6 

? 

? 

7637-8  juv. 

Tex.,  Rio  Bravo 

A.  Schott 

7640  juv. 

"     Brazos  R. 

G.  C.  Shumard 

7641  juv. 

"     El  Paso  del  Norte 

Dr.  Webb 

7642  cf  adol. 

"     Brownsville 

? 

7644  juv. 

n                a 

? 

7700  (029536) 

ad. 

"     New  Braunfels 

? 

7701  juv. 

"     Rio  Grande  del 

Norte 

G.  Wurdemann 

7747  juv. 

"     Rio  Seco 

Capt.  Pope 

stejneger:  American  soft-shell  turtles 


71 


7854  juv. 


7855  juv. 

Tex. 

,  Brownsville 

Dr.  Kennedy 

(Cotype  of 
Aspidonedes 
emoryi 
Agassiz) 

8925 

tt 

tt 

J.  C.  Merrill 

May,  1877 

10789  juv. 

tt 

San  Antonio 

C.  W.  Schuermann  June  1879 

17823  juv. 

Ark. 

,  Benton 

Jordan  &  Gilbert 

19626-8 

? 

? 

20846  9  ado!. 

Tex. 

,  Ft.  Hancock 

E.  A.  M earns 

Jan.    28,  1893 

21408  juv. 

? 

f 

26426  ad. 

Tex. 

,  Ft.  Clark 

E.  A.  Mearns 

26427  9  ad. 

it 

tt 

a 

26428-36 

u 

u 

u 

45545 

.    tt 

Boquillas 

V.  Bailey 

May  25,  1901 

46073 

It 

mouth  of  Devil's 

River 

W.  Lloyd 

Sept,  26,  1890 

46074  juv. 

II 

Eagle  Pass 

ti 

Oct.    22,  1890 

55601 

(1 

McLennan  Co. 

J.  Hurter 

1906 

66147  juv. 

La., 

Madisonville 

f 

May  29,  1886 

71627-8  adol. 

Ariz 

.,  Phoenix 

V.  Housholder 

May     1,  1926 

78515-6  juv. 

Tex. 

,  Coleto  Creek 

J.  D.  Mitchell 

Oct.,  1905 

78517 

Tex. 

,  Guadalupe  R. 

J.  D.  Mitchell 

Aug.,  1912 

83690 

tt 

Christoval 

C.  E.  Burt 

Apr.   25,  1931 

94335  d*  ad. 

tt 

near  Houston 

A.  C.  Chandler 

94336  c?  ad. 

tt 

Houston 

a 

94456-7 

tt 

Orange 

A.  H.  Wright 

Apr.    17,  1934 

95386  c5  juv. 

tt 

16 V2  mi.  S.E.  of 

Caddo  Lake 

C.  E.  Burt 

Apr.      1,  1934 

95773  pull. 

tt 

Llano  River,  Kimble 

Co. 

S.  Mulaik 

Aug.  10,  1933 

100089  cf  adol. 

La., 

near  Napoleonsvilk 

:    C.  E.  Burt 

1935 

100090  juv. 

a 

(i 

a 

a 

100380  d"  adol. 

a 

Plaquemine 

a 

June     8,  1935 

100419  <?  adol. 

a 

Spanish  Lake  near 

St, 

Gabriel 

it 

June,  1935 

103678  9 

Tex. 

,  Boquillas 

T.  Smith 

Aug.     6,  1937 

104240  9  ad. 

a 

Pecos  River  near 
Dryden 

F.  M.  Setzler 

72  bulletin:  museum  of  comparative  zoology 

Amyda  agassizii  '  (Baur) 
Plates  26-30 

1857  .—Piatypeltis  ferox  AGASSIZ,  Contr.  Nat.  Hist.  United  States,  pt.  1,  p. 

401  (part)  (not  of  Schneider). 
Aspidonectes  ferox  COKER,  North  Carolina  Geol.  Surv.,  Bull.  No.  14, 

1906,  p.  66  (South  Carolina:  Darlington  Co.,  Society  Hill,  Peedee 

River).— CORRINGTON,  Copeia,  No.  172,  Nov.  15,  1929,  p.  82 

(Congaree  Riv.,  between  Columbia  and  New  Brookland,  Lexington 

Co.,  South  Carolina). 
1888.     Piatypeltis  agassizii  BAUR,  Amer.  Natural.,  22,  p.  1121  (type  locality, 

Georgia;  type  M.C.Z.  no.  37172). 
Pelodiscus  agassizii  BAUR,  Proc.  Amer.  Philos.  Soc,  31,  July  1893, 

p.  218. 
Trionyx  agassizii  HAY,  Indiana  Geol.  17  Rep.,  1892,  p.  552;  Batr. 

Rept.  Indiana,  1893,  p.  144  (part,  U.S.N.M.  no.  8359). 
1899.     Aspidonectes  agassizii  JORDAN,  Man.  Vert.  North  Amer.,  ed.  8,  p. 

206  (part  only:  Ga.)  (emendation). 
1923.     Trionyx  spiniferus  agassizii  SIEBENROCK,  Verh.  Zool.  Bot.  Ver. 

Wien,  73,  Aug.  1923,  p.  188  (part:  Georgia). 
1939.— Trionyx  spiniferus  POPE,  Turtles  U.  S.,  Canada,  pi.  45,  figs.  98-99 

(Broad  River  near  Columbia,  Richland  Co.,  South  Carolina)  (not  of 

LeSueur). 

The  agassizii  Group 

In  outward  appearance  Amyda  agassizii  differs  very  little  from  the 
members  of  the  spinifera  group,  but  the  skull  distinguishes  it  at  once 
from  all  other  American  soft-shell  turtles.  In  practically  all  the  charac- 
ters which  differentiate  it  from  them  it  agrees  with  the  Asiatic  mem- 
bers of  the  group  represented  by  the  collective  Amyda  sinensis.  A. 
agassizii  therefore  may  be  treated  as  a  member  of  a  separate  group, 
thus  more  insistently  emphasizing  its  isolated  position  and« insuring 
the  positive  identification  of  specimens  from  its  comparatively  re- 
stricted range  in  the  United  States. 

Among  Louis  Agassiz's  collection  of  colored  drawings  of  turtles  by 
Burkhardt  (mostly  from  life)  are  several  painted  in  November  and 
December  1855,  and  inscribed  as  "Trionyx  ferox,  Ga.  Dr.  Daniel." 
They  are  excellent  pictures  of  Amyda  agassizii,  two  of  an  "adult" 
specimen  (upper  and  lower  surfaces)  and  one  (upper  side)  of  a  young 
specimen,  but  unfortunately  they  are  without  indication  of  size  and 
locality.  They  were  sent  to  Agassiz  by  Dr.  W.  B.  Daniel  from  Sa- 
vannah, Ga.  (Contr.  Nat.  Hist.  U.  S.,  1,  p.  401).    The  fact  that  they 

1  Named  for  Louis  Agassiz  to  indicate  that  the  species  was  included  by  him  in  his  account  of 
Piatypeltis  ferox. 


stejneger:  American  soft-shell  turtles  73 

came  from  the  recorded  type  locality  of  T.  ferox  evidently  influenced 
Agassiz  to  regard  these  specimens  as  topo typical  of  ferox  and  hence 
he  recorded  them  as  such. 

The  status  of  Baur's  Platypeltis  agassizii  has  never  been  fully  ex- 
plained. In  1888  (Amer.  Natural.,  vol  22,  p.  1121)  the  name  appears 
for  the  first  time  in  the  following  sentence:  "  Platypeltis  ferox  of  Agassiz 
is  not  Testudo  ferox  Schneider,  but  a  new  species,  which  may  be  called 
Platypeltis  Agassizii."  No  further  description  or  indication  is  given, 
but  evidently  reference  is  intended  to  Agassiz's  account  of  the  species 
in  his  Contribution  to  the  Natural  History  of  the  United  States,  pt.  1 , 
pp.  401-403.  A  careful  examination  of  Agassiz's  text  compared  with 
the  material  available  to  him  at  the  time  he  wrote  his  account  shows, 
that  his  Platypeltis  ferox  is  a  mixture  of  two  species  due  to  the  fact 
that  the  ranges  of  both  species  overlap  in  Georgia  and  that  the  startling 
color  pattern  of  the  very  young  specimens  of  the  true  Testudo  ferox  of 
Schneider  was  unknown  by  him.  Dr.  Baur,  in  studying  Agassiz's 
material  in  the  Museum  of  Comparative  Zoology,  designated  the 
specimen  marked  "Ferox  Ga.  No.  1"  as  the  type  of  P.  agassizii  with 
his  own  hand,  and  it  must-be  accepted  as  such.1  In  1893,  after  having 
examined  skulls  and  restudying  the  question  of  the  generic  relation- 
ships of  the  Trionychids  (Proc.  Amer.  Philos.  Soc,  vol.  31,  July  1893, 
p.  217)  he  came  to  the  conclusion  that  his  P.  agassizii  was  not  only 
specifically  but  generically  different  from  Schneider's  P.  ferox  and  re- 
ferred it  to  Fitzinger's  genus  Pelodiscus  with  several  Asiatic  species. 

In  the  meantime  Dr.  O.  P.  Hay,  assuming  that  Agassiz's  (op.  cit.) 
pi.  6,  fig.  3,  of  his  young  Platypeltis  ferox  with  the  strongly  marked  two 
black  marginal  rings  represented  the  young  P.  agassizii  (which  has  a 
similar  pattern),  applied  the  name  to  the  form  which  Agassiz  had  de- 
scribed earlier  as  P.  asper  (Indiana  Geol.  17  Rep.,  1892,  p.  552;  Batr. 
Rept.  Indiana,  1893,  Batr.  Rept.  Indiana,  1893,  p.  144).  This  con- 
fusion between  A.  agassizii  and  A.  asper  before  their  diagnostic  fea- 
tures were  well  understood  was  caused  by  the  specimen  U.S.N.M.  No. 
8359,  a  young  A.  agassizii,  which  was  alluded  to  by  Dr.  Hay  (op.  cit.)— 
though  without  giving  the  number — under  A.  agassizii  as  follows: 
"This  species  belongs  to  the  Southern  States  from  South  Carolina  to 
Texas.  A  single  specimen  has  been  forwarded  to  the  National  Museum 
from  Madison,  Ind.".  The  specimen  has  the  2  black  rings  of  the  disk 
characteristic  of  A.  agassizii  very  plainly  marked  (pi.  30,)  and  as  this 
is  also  the  normal  pattern  of  the  very  young  A.  asper,  Dr.  Hay  took 

1  In  a  letter  dated  Feb.  13,  93  to  Stejneger,  he  wrote:  "The  only  Pelodiscus  agassizii  which  I 
have  seen  is  at  Cambridge  Mus.  (the  type)." 


74 


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stejneger:  American  soft-shell  turtles 


75 


for  granted  that  the  agassizii  and  asper  were  synonymous,  an  error 
followed  by  many  subsequent  authors. 

The  fact  is  that  the  locality  "Madison,  Ind."  attributed  to  specimen 
No.  S359  in  the  original  book  of  entry  on  June  25,  1875,  is  erroneous. 
There  is  no  record  as  to  the  donor  or  collector  of  the  specimen  and  it 
was  entered  on  the  book  as  part  of  an  accumulation  of  miscellaneous 
material.  It  is  catalogued  as  Trionyx  ferox  and  there  is  no  original 
label  attached  to  the  specimen,  which  only  bears  the  tin-tag  8359,  at 
a  time  when  the  Museum  had  started  a  campaign  to  collect  living 
reptiles  and  amphibians  to  serve  as  models  for  the  series  of  painted 
casts  in  the  exhibition  series  of  the  North  American  vertebrate  fauna. 
The  tin-tagging  and  uncritical  re-entry  of  many  old  specimens  the 
record  of  which  was  lost  has  been  the  source  of  many  errors.  (Note 
reference  to  this  specimen  by  Cahn  in  his  Turtles  of  Illinois,  1937,  p. 
200,  under  Amyda  ferox). 


Geographical  Distribution 
Rivers  of  Georgia  and  South  Carolina;  North  into  southern  North 


Carolina. 


List  of  specimens  in  U.  S.  National  Museum 


8359  juv. 
8708  9  adol. 
029034   9 
30822  juv. 
51981  (M.C.Z.  1598) 

cf  adol. 
66859  9  adol. 
71681    9  adol. 

91282-3  ad.   9 , 

juv.  d" 
91310  9  adol. 

91311-2  juv. 


Ind.,  Madison  ?? 
Ga.,  Milledgeville 
Ga.  ? 

Ga.,  Baker  Co. 
) 

Ga.,  Savannah 
Ga.,  Augusta 
S.C.,  Greenwood 


T.  H.  Bean 
R.  Hessel 
Brimley  Bros. 

W.  B.  Daniel 
S.  F.  Hildebrand 
Dr.  Barrett, 
A.  L.  Pickens 


July,  1876 
June  7,  1902 


91491  juv. 

91533 

92521-3  J*  adol. 

92583  (M.C.Z.)  adol.  Ga. 

92584  (M.C.Z.  1601) 

9  Ga.,  Savannah 


Ga.,  above  Price  Id., 

Savannah  River  E.  H.  Wood  July,  193 

S.C.,  5  mi.  W.  of  Plum 

Branch,  Savannah  R.  E.  H.  Wood 
S.C.,  5  mi.  W.  of  Plum 

Branch,  Savannah  R.  "  " 

S.C.,  Batesburg  L.  Brodie 

"    Murray  Lake  C.  E.  Burt  July  5,  1933 

"    Parks ville  K.  McNeill  1933 


PLATES 


PLATE  1 


Stejnegeh — Soft-Shell  Turtles 


PLATE  1 

Skull  of  Amy  da  (spinifera) 

Fig.  1.  View  from  above.  Fig.  2.  View  from  below.  Fig.  3.  View  from  left 
side,  including  mandible.  Fig.  4.  View  of  mandible  from  above.  Fig.  5.  An- 
terior plastral  outlines  of  Amyda  ferox  U.S.N.M.  60496  (upper);  Amyda 
emoryi  U.S.N.M.  94456  (middle);  Amyda  spinifera  U.S.N.M.  101386  (lower). 


alv.  mb. 

Alveolar  surface  of  man- 

pm. 

premaxillary  (intermaxil 

dible 

lary) 

alv.  mx. 

Alveolar  surface  of  max- 

pof. 

postfrontal  (postorbital) 

illa 

prf. 

prefrontal 

art. 

articular 

pro. 

prootic  (otosphenoid) 

boc. 

basioccipital 

pt. 

pterygoid 

bsp. 

basisphenoid  (para- 

qj- 

quadratojugal  (para- 

sphenoid) 

quadratum) 

cho. 

choana 

qu. 

quadrate 

cond. 

occipital  condyle 

qu.  art. 

articulation  of  quadrat* 

cor. 

coronoid 

with  lower  jaw 

den. 

dentary 

s.  ang. 

supraangular 

exoc. 

exoccipital 

soc. 

supraoccipital 

fr. 

frontal 

splen. 

splenial 

int.  max. 

for.  intermaxillary   foramen 

sq. 

squamosal 

J"- 

j«gal 

sym. 

symphysial 

mx. 

maxillary 

temp,  fossa 

temporal  fossa  (inter 

orb. 

orbit 

temporal  foramen) 

pa. 

parietal 

tymp.  cav. 

tympanic  cavity 

pal. 

palatine 

vom. 

vomer 

paoc. 

paroccipital  (opisthotic) 

BULL.   MUS    COMP.  ZOOL 


-temp 
fossa. 


Stejneger.    Soft-Shell  Turtles.    Plate  1. 

pm. 

~mt  max  for 


alv.  mx 


temp, 
fossa 


Fig.  1 


Fig.  2 


pro 
pot    ju.        Pil-MJ- 


sopaoc 


s.ar> 


Fig.  4 


Fig. 


I 

PLATE  2 


Stejneger — Soft-Shell  Turtles 


PLATE  2 
Amyda  mutica  9,  Fairport,  Iowa.   U.S.N.M.  53521. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  2. 


■  «  • 

V      < 

,   w 
A   ¥  A 


<H- 


3Sri 


PLATE  3 


Stejneger — Soft-Shell  Turtles 


PLATE  3 

Upper  view  of  Amyda  mutica,  U.S.N.M.  95186  from  Medicine  Lodge  River, 
near  Lake  City,  Kansas.   Also  left  side  of  head. 


BULL     MUS    COMP    ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  3. 


PLATE  4 


Stejneger — Soft-Shell  Turtles 


PLATE  4 

Lower  view  of  Amyda  mutica,  U.S.N.M.  95186  from  Medicine  Lodge  River, 
near  Lake  City,  Kansas. 


BULL.    MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  4. 


PLATE  5 


Stejneger — Soft-Shell  Turtles 


PLATE  5 

Skull  of  type  of  Amyda  ferox  in  British  Museum,  from  Georgia. 


BULL     MUS    COMP.  ZOOL. 


Stejneqer.    Soft-Shell  Turtles.    Plate  5. 


PLATE  6 


Stejneger — Soft-Shell  Turtles 


PLATE  6 

Skulls  of  Amydaferox  (U.S.N.M.  029475,  029459,  029464  and  029462  from 
Kissimmee,  Florida)  showing  extremes  in  the  width  of  the  alveolar  surface  and 
in  the  outline  of  the  maxilla. 


BULL.   MUS    COMP    ZOOL. 


Stejneger.    Sot-Shell  Turtles.    Plate  6. 


PLATE  7 


Stejneger — Soft-Shell  Turtles 


PLATE  7      . 

Upper  view  of  amyda  ferox,  U.S.N.M.  86828,  Tamiami  Trail  near  Birdon, 
Florida. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.     Plate  7. 


PLATE  8 


Stejneger — Soft-Shell  Turtles 


PLATE  8 

Lower  view  of  A  my  da-  ferox,  U.S.N.M.  86828,  Tamiami  Trail  near  Birdon, 
Florida. 


BULL.    MUS.  COMP.  ZOOL. 


Stejneger.    Suet-Shell  Turtles.    Plate  8. 


PLATE  9 


Stejneger — Soft-Shell  Turtles 


PLATE  9 

Amyda  ferox,  U.S.N. M.  86492  from  Tamiami  Trail  about  15  miles  from 
Miami  City,  Florida. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  9. 


\ 


mm 


PLATE  10 


Stejneger — Soft-Shell  Turtles 


PLATE  10 

Amydaferox,  U.S.N.M.  60496  from  Auburndale,  Florida. 


BULL.   MUS    COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  10. 


PLATE  11 


Stejneger — Soft-Shell  Turtles 


PLATE  11 

Upper  view  of  Amyda  spinifera,  U.S.N.M.  101386  from  Middle  Fork,  Hol- 
ston  River,  Seven  Mile  Ford,  Virginia. 


BULL.   MUS.  COMP.  ZOOL 


Stejneger.    Soft-Shell  Turtles.    Plate  11. 


i 


"   t« 


H 


'* 


I 


I? 


Bg?       ' 


PLATE  12 


Stejneger — Soft-Shell  Turtles 


PLATE  12 

Lower  view  of  Amyda  spinifera,  U.S.N.M.  101386  from  Middle  Fork,  Hol- 
ston  River,  Seven  Mile  Ford,  Virginia. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.     Soft-Shell  Turtles.    Plate  12. 


PLATE  13 


Stejneger — Soft-Shell  Turtles 


PLATE  13 

Head  of  Amyda  spinifera,  U.S.N.M.  101386  from  Middle  Fork,  Holston 
River,  Seven  Mile  Ford,  Virginia. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  13. 


PLATE  14 


Stejneger — Soft-Shell  Turtles 


PLATE  14 

Cotype  of  Amyda  nuchalis,  Mus.  Comp.  Zool.  1623  from  Cumberland  River, 
Tennessee.  Upper  view. 


BULL.    MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.     Plate  14 


^M 


zJs 


PLATE  15 


Stejneger — Soft-Shell  Turtles 


PLATE  15 

Cotype  of  Amyda  nuchalis,  Mus.  Comp.  Zool.  1623  from  Cumberland  River, 
Tennessee.   Lower  view. 


BULL     MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  15. 


PLATE  16 


Stejneqer — Soft-Shell  Turtles 


PLATE  16 

Upper  and  lower  views,  and  side  of  head  of  Amyda  spinifera  aspera.  Cotype, 
Mus.  Comp.  Zool.  1622  from  Lake  John,  Florida. 


BULL     MUS    COMP.  ZOOL 


Stejneger.    So't-Shell  Turtles.    Plate  16. 


PLATE  17 


Stejneger — Soft-Shell  Turtles 


PLATE  17 

Amyda  spinifera  aspera,  U.S.N.M.  95191  from  Canton,  Mississippi.   Upper 
view. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  17. 


PLATE  18 


Stejnegeh — Soft-Shell  Turtles 


PLATE  18 

Amyda  spinifera  aspera,  U.S.N. M.  95191  from  Canton,  Mississippi.   Lower 
view. 


BULL     MUS    COMP.  ZOOL. 


Stejneqer.    Soft-Shell  Turtles.    Plate  18. 


PLATE  19 


Stejneger — Soft-Shell  Turtles 


PLATE  19 

a.  Amyda  ferox  juv.,  U.S.N.M.  61087  from  Auburndale,  Florida,  b.  Amy  da 
ferox  juv.,  U.S.N.M.  84603  from  Chesser's  Island,  Okefinokee  Swamp,  Georgia, 
c.  Amyda  aspera,  M.C.Z.  1608A  from  Mobile,  Alabama. 


BULL.  MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  19. 


I 

r9H 


PLATE  20 


Stejneqer — Soft-Shell  Turtles 


PLATE.  20 

Agassiz  drawing  of  Amyda  emoryi  (upper  view)  which  corresponds  fairly 
closely  with  the  type  (M.C.Z.  1910)  collected  at  Brownsville,  Texas  by  Col. 
Emory. 


BULL     MUS.  COMP.  ZOOL 


Stejneger.    Soft-Shell  Turtles.    Plate  20. 


^:$itik 


.y  i  \  >  - 


V 

V 


I 


/ 


/ 


\ 


PLATE  21 


Stbjneger — Soft-Shell  Turtles 


PLATE  21 

Agassiz  drawing  of  Amyda  emoryi  (lower  view)  which  corresponds  fairly 
closely  with  the  type  (M.C.Z.  1910)  collected  at  Brownsville,  Texas  by  Col. 
Emory. 


BULL.    MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  21. 


w 


m 


■&•>•■ 


PLATE  22 


Stejneger — Soft-Shell  Turtles 


PLATE  22 

Skull  of  Amyda  emoryi,  U.S.N.M.  78517,  collected  at  Guadalupe  River, 
Victoria  County,  Texas  by  J.  D.  Mitchell. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  22. 


fi 


Aft 


^ 


PLATE  23 


Stejneger — Soft-Shell  Turtles 


PLATE  23 

Upper  view  of  Amyda  emoryi,  U.S.N.M.  94336  from  near  Houston,  Texas. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  23. 


PLATE  24 


Stejneger — Soft-Shell  Turtles 


PLATE  24 
Lower  view  of  Amyda  emoryi,  U.S.N. M.  94336  from  near  Houston,  Texas. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  24. 


k 


1 


PLATE  25 


Stejneger — Soft-Shell  Turtles 


PLATE  25 
Head  of  Amyda  emoryi,  U.S.N.M.  94336  from  near  Houston,  Texas. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  25. 


^ 


\ 


JL-.)k 


PLATE  26 


Stejneger — Soft-Shell  Turtles 


PLATE  26 

Amyda  agassizii  Juv.,  U.S.N.M.  8359,  wrongly   recorded  from    Madison, 
Indiana.   Upper  view. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  26. 


i'l  • 


XSJii- 


^flt&'i    ° 


o 


o 


Q       0 


r 


• 


\ 


/ 


PLATE  27 


Stejneger — Soft-Shell  Turtles 


PLATE  27 

Upper  view  of  Amyda  agassizii,  U.S.N.M.  92521  from  Parksville,  South 
Carolina. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  27. 


A 


■   > 


\ 


1  i 


# 


hi, 


PLATE  28 


Stejneqer— Soft-Shell  Turtles 


PLATE  28 

Lower  view  of  Amyda  agassizii,  U.S.N.M.   92521  from  Parksville,  South 
Carolina. 


BULL     MUS.  COMP    ZOOL 


Stejneger.    Soft-Shell  Turtles.    Plate  28. 


PLATE  29 


Stejneger — Soft-Shell  Turtles 


PLATE  29 
Head  of  Amyda  agassizii,  U.S.N.M.  92521  from  Parksville,  South  Carolina. 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.    Plate  29. 


PLATE  30 


Stbjneger — Soft-Shell  Turtles 


PLATE  30 

Skull  of  Amyda  agassizii,  Philadelphia  Acad.  Nat.  Sci.  106  (  =  371). 


BULL.   MUS.  COMP.  ZOOL. 


Stejneger.    Soft-Shell  Turtles.     Plate  30. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  2 


CONTRIBUTION  TO  THE  ORNITHOLOGY  OF  THE 
HAWAIIAN  ISLANDS 


By  E.  H.  Bryan,  Jr.  and  J.  C.  Greenway,  Jr. 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED  FOR  THE  MUSEUM 

May,  1944 


PUBLICATIONS 
OF  THE    . 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1 
have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  2 


CONTRIBUTION  TO  THE  ORNITHOLOGY  OF  THE 
HAWAIIAN  ISLANDS 


By  E.  H.  Bryan,  Jr.  and  J.  C.  Greenway,  Jr. 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED   FOR  THE  MUSEUM 

May,  1944 


No.  2 — Contribution  to  the  Ornithology  of  the  Hawaiian  Islands  ' 
By  E.  H.  Bryan,  Jr.  and  J.  C.  Greenway,  Jr. 

FOREWORD 

For  over  a  year  prior  to  the  entrance  of  the  United  States  into  the 
war,  Mr.  E.  H.  Bryan,  Jr.,  Curator  of  Collections  at  the  B.  P.  Bishop 
Museum,  and  Mr.  J.  C.  Greenway,  Associate  Curator  of  Birds  at  the 
Museum  of  Comparative  Zoology,  collaborated  in  the  preparation  of 
a  Check-List  of  Birds  of  the  Hawaiian  Islands.  The  finished  draft  of 
the  list  was  completed  early  in  1941,  but,  due  to  the  fact  that  the 
senior  author  was  called  to  active  duty  as  an  army  reserve  officer, 
his  final  comments  were  not  received  for  inclusion  until  recently.  In 
the  meantime  the  junior  author  was  commissioned  in  the  Navy;  con- 
sequently the  final  preparation  of  the  Check-List  for  the  printer 
devolved  upon  the  undersigned.  No  introduction  had  been  prepared, 
but  I  was  so  fortunate  as  to  discover  in  the  Bryan-Greenway  corres- 
pondence an  outline  of  the  history  of  Hawaiian  ornithology  which, 
though  not  prepared  with  such  a  purpose  in  mind,  nevertheless  appears 
suitable  as  such.  A  brief  abstract  has  previously  appeared  in  the 
Proceedings  Hawaiian  Academy  of  Science  for  1935  (Special  Publica- 
tion no.  26,  Bernice  P.  Bishop  Museum)  p.  5-6. 

Shortly  before  being  ordered  to  active  duty  Greenway  had  incor- 
porated his  studies  of  the  Hawaiian  drepanids  in  a  rearrangement  of 
the  genera  of  this  most  interesting  family;  with  the  publication  of  this 
Check-List,  the  opportunity  to  publish  this  account  seems  opportune. 

Ornithologists  all  wish  for  the  safe  and  speedy  return  to  their  chosen 
fields  of  Capt.  Edwin  H.  Bryan,  Jr.  A. U.S.  and  Lieut.  James  C. 
Greenway,  Jr.  U.S.N.R. 


James  L.  Peters 


Cambridge,  Mass. 
23  June  1943 


1  Published  with  the  aid  of  a  special  gift  from  Mr.  G.  R.  Agassiz. 


80  bulletin:  museum  of  comparative  zoology 

HAWAIIAN  BIRDS1 

By  E.  H.  Bryan,  Jr. 

Curator  of  Collections,  B.  P.  Bishop  Museum2 

In  1778,  when  the  third  voyage  of  Captain  Cook  made  contact 
between  Hawaii  and  the  so-called  civilized  world,  there  lived  in  Hawaii 
about  100  different  kinds  of  birds.  Most  of  these  were  well  known  to 
the  Hawaiians,  each  kind  being  called  by  a  native  name.  A  few  species 
were  eaten;  the  feathers  of  several  kinds  were  used  to  decorate  the 
kahili  or  royal  standards  of  the  kings  and  high  chiefs ;  feathers  of  the 
mamo,  oo,  iiwi,  and  to  a  less  extent  those  of  the  apapane  and  o-u, 
were  tied  in  decorative  patterns  upon  fine  mesh  network  of  olona 
fiber  to  form  royal  uniforms  and  ceremonial  robes;  and  in  one  way  or 
another  many  found  a  place  in  native  life  and  lore. 

Although  Captain  Cook  did  not  live  to  see  his  native  land  again, 
and  although  there  was  no  trained  naturalist  with  this  third  voyage, 
his  companions  seem  to  have  procured  specimens  of  about  sixteen 
species  of  Hawaiian  birds,  and  to  have  carried  them  safely  back  to 
ornithologists  in  Europe.  These  first  native  Hawaiian  species  to  be 
made  known  to  science  included  the  iiwi,  mamo,  apapane,  amakihi, 
akepa,  akialoa,  o-u,  oo,  thrush,  elepaio,  rail,  crow  and  migrant  golden 
plover.  A  specimen  of  the  oo-aa,  was  also  taken,  but  this  was  not 
recognized  as  distinct  from  the  oo,  until  1855. 

The  iiwi  has  the  distinction  of  being  the  first  native  Hawaiian 
species  of  bird  to  be  technically  described.  Barthold  Lohmann,  who 
sailed  with  Cook's  last  expedition,  brought  specimens  of  it  to  Cassel, 
Germany,  where  it  was  described  by  Georg  Forster,  in  1780  as  Certkia 
coccinea.  During  the  next  four  years  John  Latham  gave  brief  descrip- 
tions and  popular  names  to  nine  other  species  in  his  General  Synopsis 
of  Birds.  In  1788,  Gmelin  gave  scientific  names  to  eleven,  including 
these,  largely  on  the  basis  of  Latham's  notes.  Six  species  can  be  recog- 
nized from  the  names  and  descriptions  given  by  James  King  in  his 
account  of  Cook's  voyage. 

Thus  did  Hawaiian  ornithology  get  away  to  a  good  start,  only  to 
sink  into  quiescent  repose  until  toward  the  end  of  the  19th  century. 
This  came  about  partly  through  neglect,  but  to  quite  a  large  extent 
because  of  a  series  of  unfortunate  circumstances. 

1  Address  of  the  retiring  president,  Hawaiian  Academy  of  Science,  May  18,  1935. 

2  Published  by  permission  of  the  Director,  B.  P.  Bishop  Museum. 


BRYAN   AND  GREE.WVAY:  HAWAIIAN'  BIRDS  81 

In  the  first  place,  the  types  of  these  early  species,  which  had  been 
placed  in  the  British  Museum,  by  some  mischance  became  lost  or 
destroyed.  This  led  to  considerable  confusion  in  later  technical  sum- 
maries, catalogs  and  lists,  because  of  the  lack  of  authentic  specimens 
to  which  to  refer.  In  the  second  place,  the  naturalists  with  Vancouver, 
Kotzebue,  and  other  early  expeditions  which  touched  Hawaii,  seem 
to  have  either  quite  neglected  the  interesting  bird  life,  or  else  to  have 
made  collections  and  observations  which  resulted  in  no  publications 
for  the  advancement  of  science. 

In  1825,  H.  M.  S.  'Blonde',  under  command  of  George  Anson, 
seventh  Lord  Byron,  carried  back  to  Hawaii  the  bodies  of  Kame- 
hameha  II  and  his  queen  who  had  died  in  England.  On  board  was 
Chaplain  Richard  R.  Bloxam  and  his  brother  Andrew,  an  enthusiastic 
young  naturalist.  Ornithologists  of  the  day  hoped  that  young  Andrew 
Bloxam  might  get  some  of  the  curious  Hawaiian  birds  and  produce 
an  interesting  publication  about  them.  He  apparently  got  very  little 
official  encouragement,  in  spite  of  which  he  obtained  specimens  of  nine 
species  of  land  birds  on  Oahu,  including  the  now  extinct  thrush.  He 
worked  hard  over  his  specimens,  and  placed  them,  all  properly  labelled, 
at  the  disposal  of  the  Lords  of  the  Admiralty.  The  scientific  "Appendix 
to  the  Voyage  of  the  Blonde",  published  in  1826,  is  said  to  have  been 
edited  bv  a  woman  who  had  onlv  a  few  of  Bloxam's  notes  to  guide  her. 
This,  combined  with  some  poor  judgment  on  the  part  of  the  ornitho- 
logical gentlemen  of  the  British  Museum,  who  identified  the  specimens, 
made  the  results,  as  Professor  Alfred  Newton  put  it1  "a  disgrace  to 
all  concerned,  since,  so  far  from  advancing  the  knowledge  of  the 
subject,  it  introduced  so  much  confusion  as  to  mislead  many  sub- 
sequent writers,"  especially  in  the  absence  of  the  specimens,  which 
disappeared  not  long  after. 

A  few  years  later  another  good  opportunity  to  advance  Hawaiian 
ornithology  was  missed.  Dr.  J.  K.  Townsend,  the  American  traveller, 
and  the  well-known  naturalist,  Thomas  Nuttall,  made  a  trip  together 
to  Hawaii.  Arriving  in  January,  1836,  they  spent  three  months  col- 
lecting on  Oahu  and  Kauai.  Townsend,  returning  at  the  end  of  the 
year,  found  the  Prussian  naturalist,  Herr  Deppe,  at  Honolulu,  and 
with  him  spent  a  few  months  in  the  pursuit  of  natural  history,  leaving 
Hawaii  in  March,  1837.  A  few  of  Deppe's  birds  were  described  by 
Lichtenstein,  and  most  of  Townsend's  specimens  have  been  carefully 
preserved,  the  bulk  of  them  in  the  Academy  of  Natural  Sciences, 

1  Alfred  Newton,  Ornithology  of  the  Sandwich  Islands,  Nature,  45,  p.  466,  1892. 


82  bulletin:  museum  of  comparative  zoology 

Philadelphia.  Had  Townsend  only  published  a  list  of  his  species, 
and  had  both  men  but  made  a  scientific  record  of  their  observations, 
the  knowledge  of  Hawaiian  bird  life  would  have  been  greatly  advanced. 

In  the  course  of  a  six  months'  stay  on  Hawaii  in  1840,  the  enthus- 
iastic collectors,  Peale  and  Pickering,  of  the  United  States  Exploring 
Expedition,  obtained  a  large  collection  of  birds.  Most  of  these  speci- 
mens were  lost  in  the  shipwreck  of  the  "Peacock,"  one  of  the  ships  of 
Commodore  Wilkes'  squadron.  Still  another  misfortune  occurred  in 
1848.  Peale's  report  on  the  birds  of  the  group  was  just  off  the  press, 
and  only  a  few  copies  had  been  distributed,  when  the  entire  stock  was 
destroyed  by  a  fire.  It  is  the  opinion  of  some  ornithologists  that  John 
Cassin's  new  edition  of  this  report,  published  ten  years  later,  was  no 
improvement  on  Peale's  original  work. 

In  1852  Dr.  Hartlaub  wrote  a  review  of  Peale's  lost  work,  and  later 
he  compiled  the  first  list  of  Hawaiian  birds,  published  in  1854.  He 
listed  but  30  species  of  birds  (of  which  five  have  not  been  accepted). 
Of  the  remaining  25,  only  16  are  land  birds,  and  only  14  are  perching 
birds. 

It  remained  for  our  own  Sanford  B.  Dole  to  produce  an  extensive 
list  of  the  Hawaiian  birds.  This  was  published  first  in  1869  in  the 
Proceedings  of  the  Boston  Society  of  Natural  History,  and  later, 
with  additions  and  corrections,  in  the  Hawaiian  Annual  for  1879. 
Judge  Dole  listed  53  species,  of  which  four  were  described  as  new. 

Although  in  the  vicinity  of  Hawaii  from  July  27  to  August  19, 1875, 
the  scientific  party  of  H.  M.  S.  'Challenger'  collected  only  24  bird 
skins,  of  which  only  one  species  was  new  to  science,  the  Hawaiian 
duck,  described  in  1878  by  P.  L.  Sclater,  as  Anas  wyviUiana. 

Revival  of  Interest 

Considerable  credit  for  a  revival  of  interest  in  Hawaiian  birds 
should  go  to  Professor  Alfred  Newton,  of  Cambridge  University, 
from  whose  interesting  account  of  early  Hawaiian  ornithology  some 
of  the  foregoing  has  been  condensed.  Because  of  Dr.  Newton's  enthu- 
siasm, two  young  Cambridge  naturalists  undertook  collecting  which 
led  to  magnificent  contributions  to  Hawaiian  ornithology. 

The  first  of  these  was  Scott  B.  Wilson  who  went  to  Hawaii  and 
collected  birds  at  Professor  Newton's  request.  Leaving  Liverpool  on 
February  24,  1887,  he  arrived  in  Honolulu  on  April  8,  having  made 
a  brief  visit  at  Washington,  D.  C,  with  Dr.  Leonhard  Stejneger, 
who  had  done  considerable  work  on  Hawaiian  birds,  especially  those 


BRYAN*  AND  GKKKNWAY:  HAWAIIAN    BIRDS  83 

sent  to  him  at  the  U.  S.  National  Museum  by  Valdamar  Knudsen  of 
Kauai.  Wilson  stayed  on  the  islands  until  the  close  of  1888,  taking 
back  with  him  to  England  a  large  and  valuable  collection  of  bird 
specimens,  rich  in  new  species.  This  collection  formed  the  basis  for 
the  beautifully  illustrated  "Aves  Hawaiienses,"  issued  in  sections 
from  1890  to  1899.  Wilson  again  visited  Hawaii,  in  1896,  without 
adding  much  to  scientific  knowledge.  Besides  collecting  skins,  he 
carefully  preserved  whole  specimens  of  the  birds  in  alcohol.  Upon 
these  specimens  Dr.  Hans  Gadow  based  anatomical  studies  which 
led  to  some  new  and  startling  conclusions  as  to  the  relationships  of 
many  Hawaiian  species. 

Meanwhile,  the  Hon.  Walter  Rothschild,  the  other  naturalist 
interested  by  Professor  Newton  at  Cambridge,  sent  his  collector, 
Henry  Palmer,  to  the  islands  to  procure  specimens  and  data.  Arriving 
in  December,  1890,  Palmer  collected  on  nearly  all  the  main  islands 
of  the  group  in  company  with  George  C.  Munro,  and  also  made  a  trip 
to  Laysan  and  Midway  Islands,  stopping  at,  or  at  least  sighting, 
most  of  the  small  rocky  islets  and  reefs  en  route.  By  August,  1893, 
when  he  left  Honolulu,  Palmer  had  collected  1832  bird  specimens, 
including  all  but  seven  of  the  species  previously  known,  and  in  addi- 
tion fifteen  which  Rothschild  described  as  newT  to  science. 

Rothschild  produced  the  elaborate  "Avifauna  of  Laysan  and  the 
neighbouring  islands  with  a  complete  history  to  date  of  the  birds  of 
the  Hawaiian  possessions,"  published  in  London  in  three  parts,  which 
appeared  August,  1893,  November  1893,  and  December  1900.  These 
contain  over  300  large,  well  printed,  folio  pages,  and  numerous  photo- 
graphs and  artistic  colored  plates  drawn  by  Keulemans  and  Frohawk. 
The  work  enumerates  116  species  of  birds,  giving  valuable  notes  on 
food,  habits,  eggs,  nests,  and  distribution,  as  w7ell  as  descriptions. 

In  1898  Dr.  H.  Schauinsland  spent  three  months  on  Laysan,  and 
in  his  fascinating  little  book,  printed  in  Bremen  in  1899,  he  lists  the 
birds  which  he  found  there.  In  1900  he  also  furnished  notes  on  a  few 
birds  of  Molokai. 

In  1892  R.  C.  L.  Perkins  came  to  Hawaii  under  the  joint  auspices 
of  the  Royal  Society  of  London  and  the  British  Association  for  the 
Advancement  of  Science,  writh  financial  assistance  from  Bernice  P. 
Bishop  Museum.  For  a  decade  he  made  extensive  collections  of 
Hawaiian  birds,  as  well  as  of  insects  and  other  land  animals.  His 
section  of  the  'Fauna  Hawaiiensis'  on  the  birds  was  published  Novem- 
ber 1903.  It  gives  an  interesting  and  valuable  summary  of  their  dis- 
tribution and  relationships,  especially  of  the  native  perching  birds. 


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In  1901,  there  appeared  a  "Key  to  the  birds  of  the  Hawaiian  group" 
by  William  Alanson  Bryan,  followed  by  a  number  of  other  lists  and 
notes  on  the  birds  of  different  islands  of  the  group,  written  by  him 
and  by  Alvin  Seale,  published  by  Bernice  P.  Bishop  Museum.  Also, 
in  1902-3,  there  was  printed  "A  complete  list  of  the  birds  of  the 
Hawaiian  possessions,  with  notes  on  their  habits,"  by  H.  W.  Henshaw. 
All  of  these  publications  have  helped  to  complete  and  bring  up  to  date 
our  knowledge  of  Hawaiian  ornithology. 

By  1930  the  large  number  of  birds  which  had  been  imported  from 
many  parts  of  the  world  prompted  the  production  of  a  report  by 
E.  L.  Caum  on  the  "Exotic  birds  of  Hawaii,"  published  by  B.  P. 
Bishop  Museum  in  1933. 

Recent  articles  on  Hawaiian  birds  include: — a  report  on  sea  bird 
conditions  on  Laysan  in  1911  by  Homer  Dill  and  W.  A.  Bryan;  four 
chapters  in  W.  A.  Bryan's  Natural  History  of  Hawaii;  a  popular 
account  of  bird  life  in  the  Hawaiian  Islands  Bird  Reservation  by 
Alexander  Wetmore;  four  chapters  in  Hawaiian  Nature  Notes  by  E.  H. 
Bryan,  Jr. ;  and  various  articles  in  journals  and  the  daily  press. 


Summary  of  Hawaiian  Birds 

With  this  brief  historical  background,  let  us  consider  the  different 
groups  of  birds  which  are  found  in  Hawaii,  and  some  of  the  interesting 
facts  concerning  them.  The  list  of  the  birds  reported  as  having  been 
found  in  Hawaii,  outside  of  captivity,  both  native  and  introduced, 
now  numbers  232  species  (see  table  1.)  Of  these,  77  species  might  be 
considered  as  endemic;  18  are  sea  birds  which  nest  regularly  in  the 
archipelago,  but  which  have  a  wider  range;  1  species,  the  night  heron, 
is  indigenous,  but  is  found  elsewhere;  8  are  regular  winter  migrants; 
34  have  been  recorded  from  time  to  time,  and  might  be  classed  as 
occasional  migrants  or  chance  arrivals;  and  94  are  immigrant  exotic 
species.  Of  these  introduced  species,  53  are  probably  established,  and 
41  are  probably  not  established,  although  the  exact  status  of  several 
is  difficult  to  determine.  So  little  field  work  has  been  done  during  the 
past  30  years  that  it  is  not  wise  to  state  what  species  are  really  extinct. 

Without  reference  to  taxonomic  classification,  most  of  the  Hawaiian 
birds  may  be  divided  into  three  fairly  well  defined  groups:  (1)  the 
birds  of  the  mountain  forests;  (2)  the  sea  birds  and  migrants;  and 
(3)  the  introduced  species.  A  few  lowland  species,  such  as  the  two 
birds  of  prey,  the  black-crowned  night  heron,  the  native  duck  and 


BRYAN  AND  GRKENWAY:  HAWAIIAN  BIRDS 


85 


goose,  the  extinct  rail,  the  mud  hens,  the  stilt,  and  the  peculiar  land 
birds  of  Laysan,  now  all  but  extinct  on  that  island,  which  do  not  fall 
into  these  three  general  groups,  might  be  given  separate  consideration. 


Table  1.    NUMERICAL  SUMMARY  OF  HAWAIIAN  BIRDS 


Family 

Endemic 

Indigenous 

&  wide- 
ranging  sea 

Regular 

Migrants 

<  •(■(visional 

&  chance 

arrivals 

Tnt 
Estab. 

roduced 

Not  Estab 

Diomedeidae 

2 

Procellariidae 

3 

5 

Phaethontidae 

1 

1 

Sulidae 

3 

Phalacrocoracidae 

1 

1 

Fregatidae 
Ardeidae 

1 
1 

1 

Threskiornithidae 

1 

Phoenicopteridae 
Anatidae 

3 

3 

10 

1 

1 

2 

Aceipitridae 
Falconidae 

1 

2 

1 

Cracidae 

3 

Tetraonidae 

1 

1 

Phasianidae 

16? 

3 

Numididae 

1 

Meleagrididae 
Turnicidae 

1 
1 

Gruidae 

1  ? 

Rallidae 

5 

1 

1 

Charadriidae 

5 

6 

Recurvirostridae 

1 

Phalaropodidae 
Laridae 

6 

2 
9 

2 

Columbidae 

7 

11 

Psittacidae 

2 

5 

Strigidae 
Alcedinidae 

1 

1 

Alaudidae 

2 

1 

Corvidae 

1 

Paridae 

1 

Timaliidae 

3 

1 

Mimidae 

1  ? 

Turdidae 

6 

2 

2 

Sylviidae 

2 

1 

Museicapidae 

3 

2 

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Prionopidae 

Sturnidae 

Zosteropidae 

Drepanididae 

45 

Meliphagidae 

5 

Ploceidae 

Icteridae 

Fringillidae 

77 

19                8 

Total  Native 

104 

Total  Introduced 

* 

Grand  Total 

232. 

The  Sea  Birds 

1  ? 

1 

1 

2 

1 

1 

1 

5 

3 

34  53  41 

94 


Concerning  the  sea  birds,  found  throughout  the  Hawaiian  group, 
little  need  be  said  here.  Most  of  them  are  species  widespread  in  the 
Pacific,  and  although  scientists  argue  over  their  scientific  names  they 
are  fairly  easy  to  recognize.  They  include:  the  two  large  albatross, 
the  thieving  frigate,  three  species  of  homely  boobies,  five  kinds  of 
noisy  terns,  the  burrowing  shearwaters  and  petrels,  and  the  graceful 
white-and-red-tailed  tropic  birds.  Most  of  them  stay  well  away  from 
man's  habitations.  Not  that  they  are  afraid  of  him.  Rather,  man 
and  sea  birds  are  rivals  for  a  large  and  steady  supply  of  fresh  fish, 
without  which  an  extensive  bird  colony  could  not  exist. 

President  Theodore  Roosevelt  and  other  thoughtful  citizens  have 
helped  to  protect  our  sea  birds  by  establishing  for  them  the  Hawaiian 
Islands  Bird  Reservation  on  islands  and  reefs  to  the  northwest  of 
Kauai.  Here  they  are  not  only  kept  from  molestation,  but  also  from 
time  to  time  efforts  are  made  to  keep  their  rather  poor  environment 
from  becoming  any  worse,  through  occasional  visits  to  kill  off  rabbits 
and  plant  more  vegetation. 

Migratory  Birds 

Each  year  there  come  to  Hawaii  large  numbers  of  tourists,  concern- 
ing whom  no  record  is  kept  by  our  efficient  Tourist  Bureau.  They 
spend  no  money  here,  but  they  do  a  very  considerable  amount  of  good. 
They  are  the  migratory  birds.  Arriving  from  a  more  than  2,000  mile 
flight,  thin  and  hungry,  they  alight  on  our  beaches,  fields  and  pastures 
and  make  short  work  of  great  numbers  of  caterpillars,  grasshoppers 


BRYAN  AND  GBEENWA1  :  HAWAIIAN  BIRDS  87 

and  other  insects,  much  to  the  benefit  of  agriculture.  They  arrive  and 
depart  with  great  regularity,  coming  in  the  late  summer  or  early  fall 
and  leaving  again  in  the  spring,  to  return  to  nest  in  their  cold  northern 
home.  Included  among  these  gentle  and  helpful  visitors,  which  should 
be  given  a  better  welcome  than  the  hunters'  gun,  are  the  golden 
plover,  the  turnstone„  wandering  tatler,  sanderling,  curlew,  various 
wild  ducks,  and  several  others. 

The  native  birds  of  Hawaii's  lowlands,  marshes,  ponds,  valleys, 
and  grassy  slopes  are  becoming  very  rare.  The  flightless  rail  is  gone; 
the  duck  and  goose  are  making  a  last  valiant  stand  against  extermina- 
tion; the  mud  hens,  stilts,  and  night  heron  seem  to  be  holding  their 
own.  One  still  sees  Pueo,  the  native  owl,  winging  his  low  flight  above 
the  tops  of  guava  bushes  and  across  grassy  hillsides  at  sunset,  in  search 
of  rats  and  lizards.  But  Io,  the  native  hawk,  is  scarce  in  Hawaii,  due 
to  the  mistaken  idea  that  it  was  an  enemy  to  the  poultrymen. 


Birds  of  the  Mountain  Forests 

The  birds  of  the  mountain  forests  are  all  native  perching  birds  and 
include  50  to  55  endemic  species  of  that  order.  Twenty-three  of  the 
twenty-four  genera  in  this  group  are  also  endemic,  only  Corpus  being 
found  elsewhere  in  the  world.  These  undoubtedly  make  up  the  most 
interesting  part  of  the  Hawaiian  bird  fauna.  Whence  did  they  come? 
What  were  their  ancestral  relationships?  Why  are  there  so  many 
genera  and  species,  even  one  whole  family,  found  nowhere  else  in  the 
world?  Why  are  they  so  rapidly  becoming  extinct?  These  are  ques- 
tions which  interest  not  only  the  ornithologist,  but  also  the  zoologist, 
the  biologist,  the  conservation  expert,  and  the  student  of  evolution. 
I  can  only  try  to  hint  at  the  answers  to  some  of  these  questions. 

These  native  perching  birds,  found  in  the  forests  of  the  main  islands 
of  the  group,  include:  the  crow,  found  only  in  the  Kona  and  Kau 
districts  of  Hawaii;  six  different  species  of  thrushes,  five  of  which 
(found  on  five  different  islands)  are  closely  related,  and  one,  on  Kauai, 
somewhat  different  in  form,  food,  and  habits;  three  closely  related 
species  of  elepaio  or  flycatcher,  on  three  different  islands;  four  species 
of  oo,  on  four  different  islands,  and  one  other  very  distinct  honey-eater 
of  Hawaii;  and  between  40  and  45  different  species  of  native  "honey 
creepers,"  belonging  to  a  family  found  nowhere  else  in  the  world. 
I  give  this  range  of  number  for  the  drepanids  because  various  authori- 
ties differ  in  their  concepts  of  specific  differences. 


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Native  Hawaiian  Honey  Creepers 

The  drepanids  themselves  are  worthy  of  extensive  study.  There 
are,  let  us  say,  forty  species,  representing  eighteen  genera.  These 
have  very  diverse  appearance;  some  are  black  and  golden,  some  red 
and  black,  some  green  and  yellow;  some  have  shprt  bills  for  picking  up 
insects,  some  long,  slender  bills  for  sipping  nectar  from  the  base  of 
the  long-tubed  blossoms  of  the  native  lobelias,  and  some  have  heavy, 
almost  finch-like  or  parrot-like  beaks  for  cracking  seeds.  Only  three 
species,  the  iiwi,  apapane,  and  o-u,  are  found  on  more  than  two  islands. 
Yet  it  is  believed,  that  all  are  the  descendents  of  a  common  ancestor, 
or  at  most  two  ancestral  stocks. 

The  careful  anatomical  studies  of  Dr.  Hans  Gadow  suggest  that 
their  nearest  relatives  are  the  Neotropical  and  Central  American 
Coerebidae.  Peter  P.  Suskin  (International  Ornith.  Kongress  1926, 
Verhandl.,  VI,  pp.  375-381,  1929)  suggests  descent  from  Fringillidae 
of  the  Malaysian  region.  We  can,  perhaps,  imagine  that  many  thou- 
sands of  years  ago  there  reached  Hawaii  from  far  away  tropical  America 
or  southeastern  Asia  a  little  group  of  these  birds.  A  pair  of  them, 
at  least,  survived,  and  establishing  themselves,  became  the  common 
ancestors  of  our  interesting  native  family.  We  can  but  suppose  that 
their  descendents,  becoming  isolated  on  different  islands  of  the  group, 
with  different  kinds  of  environment  and  different  sources  of  food 
supply,  developed  into  the  diverse  species  which  we  find  today,  or 
rather,  in  many  cases, — yesterday,  for  a  number  are  very  rare,  even 
extinct. 

Origin  of  Hawaiian  Bird  Groups 

The  same  explanation  as  to  origin  might  be  offered  for  other  groups 
of  native  perching  birds.  For  example,  one  ancestral  kind  of  immigrant 
flycatcher,  arriving  not  so  many  centuries  ago,  has  given  rise  to  the 
three  species  of  elepaio,  one  each  on  Hawaii,  Oahu,  and  Kauai,  which 
look  so  very  much  alike  that  they  might  better  be  considered  only 
geographic  races,  rather  than  different  species.  One  immigrant  an- 
cestral form  may  have  been  the  progenitor  of  the  four  kinds  of  oo, 
the  Oahu  species  of  which  has  not  been  seen  alive  since  1837;  another 
was  the  ancestor  of  Chaetoptila,  a  Hawaii  honey-eater,  now  probably 
extinct.  The  thrushes  descended  from  two  ancestral  stocks;  one 
ancestor  giving  rise  to  one  species  each  of  A-Maui  on  Kauai,  Oahu, 
Molokai,  Lanai,  and  Hawaii,  and  the  little,  weevil-eating  Puaichi 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  89 

having  descended  from  another.  Thus,  all  of  the  native  Hawaiian 
perching  birds  might  have  developed,  as  Dr.  Perkins  has  pointed  out1 
"from  at  least  six  and  not  more  than  seven  successfully  immigrant 
species." 

It  would  make  a  fascinating  study,  by  tracing  out  the  distribution 
of  the  nearest  relatives  of  these  various  groups  of  birds,  to  try  to 
determine  whence  they  came;  and,  by  studying  their  degree  of  develop- 
ment and  evolution,  as  well  as  their  local  distribution,  to  attempt  to 
guess  how  long  they  have  been  in  Hawaii.  One  interesting  point 
is  the  fact  that,  although  the  distances  between  the  Hawaiian  Islands 
are  not  very  great,  and  although  the  native  forest  birds  have  fairly 
good  powers  of  flight,  their  distribution  is  so  definitely  limited  and 
restricted. 

Why  Native  Species  are  Becoming  Extinct 

One  hears  various  explanations  as  to  why  the  native  species  of 
Hawaiian  birds  are  so  rapidly  becoming  rare  or  extinct.  We  are  told 
by  those  who  have  not  made  a  study  of  the  subject  that  they  were 
killed  off  by  the  native  feather-gatherers  or  chased  away  by  the  minah 
birds,  or  their  eggs  destroyed  by  the  mongoose.  These,  in  certain 
instances,  may  have  been  contributing  causes,  but  they  have  certainly 
not  been  the  basic  reason  for  the  changes  which  have  come  over  the 
native  forest  life.  The  main  reason  may  be  stated  in  these  few  words: 
Nature  established  a  balance  in  Hawaii,  and  Nature's  balance  has 
been  upset.  What  is  this  "Balanee  in  Nature"  and  what  has  caused 
its  upset? 

Long  ago,  before  the  coming  of  man  with  his  ships,  new  plants  and 
animals  arrived  from  foreign  parts  and  established  themselves  in 
Hawaii  only  at  long,  infrequent  intervals  of  time.  This  was  because 
of  Hawaii's  extreme  isolation — more  than  two  thousand  miles  from 
the  regions  from  which  the  ancestors  of  most  of  the  native  species 
came.  The  kinds  which  managed  to  arrive  and  gain  a  foothold  had  to 
adjust  themselves  to  their  new  home.  If  they  were  in  harmony  with 
their  new  environment  they  survived;  if  they  were  not  in  harmony, 
they  either  became  adjusted  or  modified  until  they  were,  or  else  they 
perished.  There  have  been  many  types  of  environment  in  HawTaii: 
bare  lava  flows,  hot  dry  deserts,  fertile  valleys,  marshy  lowlands, 
grassy   hills,   wet  forests,   summit   bogs,   bare,   snow-capped  peaks. 

1  R.  C.  L.  Perkins,  Fauna  Hawaiiensis,  vol.  I,  pt.  iv,  p.  369,  1903. 


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Each  offered  a  habitat  for  a  different  type  of  life,  or  a  reason  for 
modification  or  adjustment.  In  each  different  environment  a  balance 
was  established,  with  the  ironclad  policy,  "fit  in  or  perish."  If  a 
species  became  over-abundant  it  was  brought  back  to  a  proper  numer- 
ical balance  either  through  lack  of  food  supply,  or  by  the  activity  of 
its  enemies,  which  took  advantage  of  its  abundance  to  multiply  them- 
selves, only  in  turn  to  be  starved  or  eaten  back  to  normal. 


Upset  of  Nature's  Balance 

Then  into  this  nicely  balanced  island  world  came  man,  with  his 
foreign  plants  and  animals,  his  thoughtless  or  ruthless  waste,  his 
clearing  and  burning.  The  nice  balance  of  Nature  was  upset.  Cattle, 
sheep,  pigs,  and  goats  ran  wild  through  the  forests,  so  damaging  the 
undergrowth  that  it  no  longer  protected  the  roots  of  the  growing  trees. 
Rain  eroded  away  the  soil;  seeds  could  no  longer  germinate;  and  both 
forest  trees  and  undergrowth  died  off.  Their  places  were  quickly 
taken  by  foreign  plants  which  no  longer  furnished  food  or  protection 
to  native  insects  and  birds.  Whole  associated  groups,  interdependent 
upon  each  other,  were  disassociated.  Foreign  birds  were  imported, 
bringing  with  them  diseases  against  which  the  specialized  native 
birds  had  no  immunity.  With  food  and  shelter  destroyed,  and  danger 
threatening  them  in  the  lowlands,  the  native  birds  either  sought  new 
homes  in  the  depths  of  the  forests,  where  they  in  turn  upset  nicely 
adjusted  Nature,  or  else  they  became  extinct. 


Foreign  Birds 

And  now,  because  native  birds  are  scarce,  the  cry  is  for  more  and 
yet  more  foreign  birds.  Some  kinds  have  been  introduced  to  satisfy 
the  desire  of  the  hunter;  pheasants  from  the  mountains  of  Mongolia 
or  Java;  grouse  and  quail  from  northwestern  America,  China,  Japan 
and  Australia;  partridges  from  the  slopes  of  the  Himalayas;  doves 
and  pigeons  from  Panama,  India,  China,  Australia,  and  the  Malay 
Isles.  Some,  like  the  skylark  and  cardinal,  have  been  brought  because 
of  sentiment  or  to  satisfy  esthetic  taste.  Some  have  been  introduced 
for  a  more  useful  purpose,  such  as  the  horn  fly-eating  willie  wag-tail, 
the  meadow  lark,  even  the  much  maligned,  but  very  useful  minah. 
A  few  accidental  escapes  we  could  well  get  along  without. 


BRYAN'  AND  GREENWAY:  HAWAIIAN   BIRDS  91 

Fortunately  for  Hawaii,  our  Territory's  scientific-  advisers  have 
advocated  a  policy  of  rigid  protection  for  our  native  and  useful  bird 
life,  and  a  strong  stand  against  the  promiscuous  importation  of  foreign 
birds  and  animals.  This  need  not  include  birds  which  are  to  be  kept 
in  a  proper  aviary,  carefully  safeguarded  against  their  escape.  I  can 
see  much  that  is  good  in  recent  plans  for  extensive  breeding  and 
exhibition  of  showy  and  interesting  birds  in  Honolulu,  so  long  as  they 
are  not  allowed  to  go  wild.  But  I  cannot  approve  of  wholesale  intro- 
duction and  liberation,  without  adequate  study  by  experts  as  to  the 
consequences. 

You  may  say,  now  that  Nature's  balance  has  been  so  badly  upset, 
and  Hawaii's  avian  heritage  is  passing  out  of  the  picture,  what  harm 
can  a  little  more  upset  do !  But  the  birds  may  not  be  the  only  part  of 
our  natural  history  to  be  effected  by  such  introductions.  Forestry, 
agriculture,  and  all  branches  of  animal  life  are  so  interrelated  that 
one  cannot  tell,  without  considerable  study,  what  far  reaching  con- 
sequences may  result  from  one  foolish  introduction.  We  have  effective 
quarantine  laws  to  protect  Hawaii  against  insect  pests  and  plant 
diseases,  which  we  all  admit  help  tremendously  to  safeguard  our 
island  paradise.  Let  us  try  to  protect  what  bird  life  we  have  left,  and 
also  try  to  avoid  the  risk  of  further  upset  of  Nature's  balance,  just 
for  the  sake  of  adding  a  flash  of  color  or  a  chirp  of  song. 


92  bulletin:  museum  of  comparative  zoology 

'    CHECK-LIST  OF  THE  BIRDS 
OF  THE  HAWAIIAN  ISLANDS 

By  E.  H.  Bryan,  Jr.  and  J.  C.  Greenway,  Jr. 

Part  I.    NATIVE  BIRDS 

Order  PROCELLARIIFORMES 

Family  DIOMEDEIDAE— Albatrosses 

Genus  Diomedea — Albatrosses 

Diomedea  Linnaeus,  Syst.  Naturae,  ed.  10,  1,  1758,  p.  132.    Type,  by  subse- 
quent designation,  D.  exulans  Linn. 

Diomedea  nigripes  Audubon 

Diomedea  nigripes  Audubon,  Ornith.  Biog.,  5,  1839,  p.  327.    (Pacific  Ocean, 

lat.  30°  44'  N.,  long.  146°  W.) 
Black-footed  Albatross,  Brown  Gooney 
Phoebastria  nigripes  reischekia  Mathews,  Bull.  British  Ornith.  Club,  51,  1930, 

p.  29.    (New  Zealand.) 
Diomedea  chinensis  Temm.  in  Rothschild,  Avifauna  of  Laysan,   etc.,  p.  55 

(corrected  p.  292). 
Diomedea  albatrus  (chinensis)  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  xxv. 

Range.  North  Pacific,  breeding  on  islands  northwest  of  Hawaii; 
Marshall  Islands  and  Bonin  Islands. 

Recorded  in  Hawaiian  Group  from  Midway  (breeding),  Pearl  and 
Hermes  Reef  (sight),  Lisianski,  Laysan  (breeding),  French  Frigate 
Shoal  (breeding)  (sight),  Necker  (sight),  "Bird  Rock"  (Nihoa)  (sight), 
Kaula  (breeding),  Lehua  (sight)  (Caum).  Many  sight  records  at  sea. 
"This  is  the  Gooney  that  follows  in  the  wrake  of  ships  from  San  Fran- 
cisco to  Honolulu  and  Hilo"  (Henshaw). 

Nests  behind  the  beaches  on  the  periphery  of  the  islands.  A  hybrid, 
D.  nigripes  x  D.  immutabilis,  from  Midway  is  in  the  Bishop  Museum. 

Diomedea  immutabilis  Rothschild 

Diomeda  immutabilis  Rothschild,  Bull.  British  Ornith.  Club,  1,  1893,  p.  xlviii. 

(Laysan  Island.) 
Gooney,  White  Albatross,  Laysan  Island  Albatross 

Range.     Central  and  North  Pacific. 

Breeds  on  Laysan  and  Midway  Islands.  Nests  in  interior  of  the 
islands.  To  be  seen  at  sea  from  islands  off  the  coast  Lower  California 
to  the  Bonin  Islands. 


BRYAN  AND  GREENWAY  :  HAWAIIAN  BIRDS  93 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Laysan 
(breeding),  Lisianski  (breeding),  Necker  Island  (breeding),  Pearl  and 
Hermes  Reef  (breeding),  Gardner  Island,  "Bird  Island"  (Nihoa) 
(breeding).  "Apparently  common  throughout  Hawaiian  main  group, 
though  it  is  not  known  to  breed  on  any  of  them"  (Henshaw). 

Family  PROCELLARIIDAE 

Genus  Puffinus — Shearwaters 

Puffinus  Brisson,  Ornithologie,  1,  1760,  p.  56;  6,  p.  130.   Type,  by  tautonomy, 
Puffinus  Brisson  =  ProceUaria  puffinus  Briinnich. 

Puffinus  pacificus  cuneatus  Salvin 

Puffinus  cuneatus  Salvin,  Ibis,  1888,  p.  353.    (Krusenstern  Island.) 

Wedge  Tailed  Shearwater,  Uau  Kane,  Uwau 

Puffinus  pacificus  laysani  Mathews,  Birds  of  Australia,  vol.  2,  1912,  p.  83. 

(Laysan  Island.) 
Puffinus  knudseni  Stejneger,  Proceed.  U.  S.  Nat.  Museum,  11,  1888,  p.  93. 

(Kauai.) 

Range.  From  about  140°  west  latitude  (two  days  from  San  Fran- 
cisco) to  the  Bonin  Islands  and  south  to  the  Equator. 

Recorded  in  the  Hawaiian  Group  from  Midway,  Pearl  and  Hermes 
Reef  (sight,  Galstoff),  Lisianski,  Laysan  (breeding),  French  Frigate 
Shoal,  Xecker  (breeding),  "Bird  Island"  (Nihoa),  Kaula,  Lehua, 
Kauai,  Oahu  (breeding  on  Mokumanu,  Manana  and  Popoia  Islets) 
(Munro). 

Puffinus  nativitatis  Streets 

Puffinus  (Nectris)  nativitatis  Streets,  Bull.  U.  S.  Nat.  Museum,  1877,  no.  7 

p.  29.    (Christmas  Island,  Pacific  Ocean.) 
Black  Shearwater,  Christmas  Island  Shearwater 

Range.  Pacific  Ocean  from  the  vicinity  of  25°  north  latitude  south 
to  the  vicinity  of  25°  south  latitude. 

Recorded  in  the  Hawaiian  Group  from  Midway,  Laysan  (breeding), 
French  Frigate  Shoal  (breeding),  "Bird  Island"  (Nihoa)  (breeding?), 
Lisianski  (Munro  ms.),  Gardner  (sight  record  by  Dr.  Isenbeck  re- 
corded by  Kittlitz  1834),  Oahu  (Mokulua  Islet)  (sight  by  Donagho), 
Mokumanu  Islet  (breeding)  (Munro). 

Puffinus  newelli  Henshaw 
Puffinus  newelli  Henshaw,  Auk,  17,  1900,  p.  246.    (Waihee  Valley,  "Ulani"  in 

error,  Maui  Island,  see  Henshaw,  Birds  of  Hawaiian  Possessions,  p.  117, 

footnote.) 
Newell' s  Shearwater,  Ao 


94  bulletin:  museum  of  comparative  zoology 

Range.     Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  ?Niihau,  Kauai  (breeding), 
Molokai,  Maui.   We  can  find  no  records  for  Lanai. 

Genus  Pterodroma — Petrels 

Pterodroma  Bonaparte,  Comptes  Rendus  Academie  Sciences,  Paris,  42,  1856, 
p.  768.  Type,  by  subsequent  designation,  Procellaria  macroptera  A.  Smith. 

Pterodroma  phaeopygia  sandwichensis  (Ridgway) 

Oestrelala  sandwichensis   Ridgway,    Baird,    Brewer    and    Ridgway's    Water 

Birds  of  North  America,  vol.  2,  1884,  p.  395.    (Hawaiian  Islands.) 
Dark-rumped  Petrel,  Uau  or  Uwau 
Oestrelala  phaeopyga  In  Wilson  and  Evans,  Aves  Hawaiiensis. 

Range.     Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Hawaii  (breeding  on  Mauna 
Kea  and  Mauna  Loa),  Kauai  (breeding),  Molokai  (breeding)  (eggs  in 
Bishop  Museum),  Lanai  (Munro). 

Henshaw  (1902)  found  that  the  mongoose  had  driven  these  birds 
from  their  nesting  grounds  on  Hawaii. 

Pterodroma  leucoptera  hypoleuca  (Salvin) 

Oestrelata  hypoleuca  Salvin,  Ibis,  1888,  p.  359.     (Krusenstern  Island,  North 

Pacific  Ocean.) 
Salvin' s  White  Breasted  Petrel,  Bonin  Island  Petrel 

Range.  Northern  Pacific  Ocean.  Breeds  on  the  Bonin  Islands  ? 
and  Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding — eggs 
collected  by  T.  M.  Blackman),  Laysan  (breeding)  (Rothschild), 
(W.  K.  Fisher  1903),  ?  Kaula  (Caum),  Lanai  (Munro). 

Mathews  (1934)  records  Pterodroma  leucoptera  longirostris  Stejneger 
(1893)  as  the  breeding  bird  of  the  Bonin  Islands.  This  is  not  the  case. 
Birds  of  the  Bonin  Islands  differ  from  Japanese  birds  and  agree  with 
Hawaiian  birds  in  having  grayer  under  wing  coverts  and  in  their 
large  size. 

Krusenstern  Rock  probably  does  not  exist,  reported  by  Captain 
Lisianski  in  lat.  22°  15'  N.,  long.  175°  37'  W.,  it  has  been  dropped 
from  the  Pacific  Islands  Pilot  of  the  British  Admiralty  (ed.  1931) 
and  though  it  is  recorded  in  the  Hawaiian  Islands  Pilot  of  the  U.  S. 
Hydrographic  Office  (ed.  1933),  there  has  been  no  actual  report  of  a 
rock  in  this  vicinity  since  1804  when  breakers  were  reported  thirty 
miles  south  of  the  supposed  position  of  the  rock.  The  type  locality 
of  this  bird  is  therefore  doubtful. 


BRYAN  AND  GREENWAY :  HAWAIIAN  BIRDS  95 

Mathews  (1934)  records  the  breeding  range  of  this  bird  as  follows: 
"Hawaiian  Group  (Laysan,  Pearl,  Hermes  [sic],  Lisianski,  Krusenstern 
Reef,  Midway  and  Ocean' Islands  and  French  Frigate  Shoal)."  We 
cannot  find  that  this  bird  has  ever  been  recorded  as  breeding  on  any 
island  except  Laysan. 

Genus  Bulweria — Petrels 

Bulweria  Bonaparte,  Nuova  Annales  Sci.  Nat.  Bologna,  8,  1842,  p.  426.  Type, 
by  monotypy,  Procellaria  bulweri  Jardine  and  Selby. 

Bulweria  bulwerii  (Jardine  and   Selby) 

Procellaria  bulwerii  Jardine  and  Selby,  Illust.  Ornith.  2,  1828,  pi.  65.  (Ma- 
deira.) 

Soft  Nosed  Petrel,  Bulwer's  Petrel 

Bulweria  bulweri  pacifica  Mathews  and  Iredale,  Ibis,  1915,  p.  607.  (Imojima, 
Bonin  Islands.) 

Bulweria  anjinho  Heineken  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  211. 

Range.  "Breeding  on  islands  off  the  coast  of  China;  the  Bonin 
Islands,  Vulcan  Islands,  the  western  Hawaiians  and  the  Marquesas 
Islands  in  the  Pacific  Ocean;  Madeira,  the  Salvages,  Canary  and 
Cape  Verde  Islands  in  the  Atlantic"  (Peters). 

Recorded  in  the  Hawaiian  Group  from  Laysan,  French  Frigate 
Shoal  (breeding),  Necker  Island  (breeding),  Kauai,  Hilo,  Hawaii 
(Henshaw),  Oahu  (skin  in  Bishop  Museum,  G.  C.  Munro  coll.). 

W.  A.  Bryan  does  not  distinguish  between  sight  records  and  col- 
lected specimens.  His  records  are  from  Necker  and  Bird  Island 
(Xihoa). 

We  quite  agree  with  Hartert  (Novitat.  Zool.,  33,  1926,  p.  326)  that 
pacifica  cannot  be  maintained. 

Family  HYDROBATIDAE 

Genus  Oceanodroma — Petrels 

Oceanodroma  Reichenbach,  Aves  Systema  Naturae,  1852  (1853  fide  Peters) 
p.  iv.   Type,  by  original  designation,  Procellaria  furcata  Gmelin. 

Oceanodroma  castro  cryptoleucura  (Ridgway) 
Cymochorea  cryptoleucura  Ridgway,  Proceed.  U.  S.  Nat.  Museum,  4,  1882, 

p.  337.    (Kauai,  Hawaiian  Islands.) 
Hawaiian  Stormy  Petrel,  Ake-Ake  (Henshaw),  Oeoe  (W.  A.  Bryan) 

Range.     Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Midway  (Bartsch),  French 
Frigate  Shoal,  Niihau,  Kauai. 

"The  natives  report  a  small  petrel,  which  they  call  by  the  above 


96  bulletin:  museum  of  comparative  zoology 

name  (Ake-Ake),  as  common  on  the  fishing  grounds  five  or  ten  miles 
off  the  coast  of  Hawaii,  and  I  have  little  doubt  that  it  is  this  species 
..."  (Henshaw). 

OCEANODROMA  MARKHAMI  TRISTRAMI  Salvin 

Oceanodroma  tristrami  Salvin,  Catalogue  Birds  British  Museum,  25,  1896, 

p.  354.    (Sendai  Bay,  Japan.) 
Sooty  Petrel,  Tristram's  Petrel,  Fork  Tailed  Petrel. 
Oceanodroma  fuliginosa  (Gmelin)  in  Rothschild,  Avifauna  of  Laysan,  etc., 

p.  308;  W.  A.  Bryan,  Birds  of  the  Hawaiian  Group,  p.  13  (footnote); 

Henshaw,  Birds  of  the  Hawaiian  Islands,  p.  119;  Oceanodroma  tristrami 

Stejneger  in  Bartsch,  Auk,  34,  1922,  p.  486;  W.  K.  Fisher,  Bull.  U.  S.  Fish 

Commission,  23,  1923,  p.  795. 

Range.     From  Japan  south  and  west  to  the  Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Midway  (Bartsch),  Laysan 
(breeding),  ?"Bird  Island"  (Nihoa)  (W.  K.  Fisher),  Lanai  (skin  in 
Bishop  Museum,  G.  C.  Munro  coll.). 

In  our  opinion,  Salvin's  description  of  tristrami  is  identifiable  and 
Hartert's  arguments  (Vogel  palaarktischen  Fauna,  p.  1416)  have 
no  force  under  the  Rules  of  Zoological  Nomenclature.  Mathews 
records  Cymochorea  (markhami)  owstoni  Mathews  and  Iredale  1915, 
as  the  breeding  bird  of  the  Hawaiian  Islands  in  the  belief  that  the 
name  tristrami  is  not  identifiable. 


Order  PELECANIFORMES 

Family  PHAETHONTIDAE 

Genus  Phaethon — Tropic  or  Bos'n  Birds 

Phaethon  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  134.  Type,  by  sub, 
sequent  designation,  Phaethon  adherens  Linn.  (Gray,  List  Genera  Birds- 
1840,  p.  80).  For  synonymy  see  Peters,  Check-List  of  Birds  of  the  World, 
1,  p.  77. 

Phaethon  rubricauda  rothschildi  (Mathews) 

Scaeophaethon  rubricauda  rothschildi  Mathews,  Birds  Australia,  4,  1915,  p.  303. 

(Laysan  and  Niihau.) 
Red-tailed  Tropic  Bird,  Bos'n  Bird,  Koaeula 
Scaeophaethon   rubricauda   brevirostris    Mathews,    Birds   Australia,    4,    1915, 

p.  303.    (Bonin  Islands.) 
Phaethon  rubricauda  Boddaert  in  Rothschild,  Avifauna  of  Laysan,  etc.,  pp.  33, 

294,  296  and  other  works. 


BRYAN  AND  GKEKXWAY:  HAWAIIAN   BIRDS  97 

Range.  Pacific  Ocean  from  the  Bonin  and  Marianas  Islands  to  the 
Hawaiian  Islands. 

Recorded  in  the  Hawaiian.  Group  from  Midway  (breeding;  eggs  in 
Bishop  Museum,  T.  M.  Blackman  coll.),  Laysan  (breeding),  French 
Frigate  Shoal  (sight  by  W.  K.  Fisher),  Necker,  Kaula,  Lehua,  Niihau, 
Mokumanu  Islet,  Oahu  (sight,  Munro),  Lanai  (sight,  Munro). 

Wilson  and  Evans  record  this  bird  breeding  on  Kauai  and  Niihau, 
and  occurring  on  Hawaii. 

Phaethon  lepturus  dorotheae  Mathews 

Phaethon  lepturus  dorotheae  Mathews,  Austral  Avian  Rec,  2,   1913,  p.   7- 

(Queensland.) 
White   Tailed  Tropic  Bird,  Salmon  Tailed  Tropic   Bird,  Bos'n  Bird,  Koae, 

Haakoae 
Phaethon  lepturus  Daudin  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  296; 

Henshaw,  Birds  Hawaiian  Islands,  p.  114,  and  other  works. 
Phaethon  aethereus  Bloxam  (nee  Linn.)  in  Wilson  and  Evans,  Aves  Hawaiiensis, 

p.  186. 

Range.     Australia  and  certain  Pacific  islands. 

Recorded  in  the  Hawaiian  Group  from  Kauai,  Oahu,  Maui,  Molokai 
(specimens  in  Mus.  Comp.  Zool.),  Hawaii. 

Hartert  (Novitat.  Zoologicae,  32,  1925,  p.  276)  remarks  that 
dorotheae  is  probably  not  distinct  from  lepturus,  which  is  the  breeding 
bird  of  Mauritius.  This  does  not  appear  to  us  to  be  the  case.  The 
species  is  in  need  of  revision. 


Family  SULIDAE 

Genus  Sula — Boobies 

Sula  Brisson,  Ornithologie,  1,  1760,  p.  60;  6,  p.  494.    Type,  by  tautonomy, 
"Sula"  =  Sula  leucogaster  Boddaert. 

Sula  sula  rubripes  Gould 

Sula  rubripes  Gould,  Synopsis  Birds  Australia,  pt.  4,   1838,  append.,  p.  7. 

(New  South  Wales  =  Raine  Island,  N.  Queensland,  fide  Mathews.) 
Tied  Footed  Booby 

Sula  piscatrix  Linn,  in  Rothschild,  Avifauna  of  Laysan,  etc.,  pp.  27,  297. 
Sula  piscator  Linn,   in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  xxv;  Henshaw, 

Birds  Hawaiian  Islands,  p.  113;  W.  A.  Bryan,  Key,  Birds   Hawaiian 

Group,  p.  15. 


98  bulletin:  museum  of  comparative  zoology 

Range.  "Islands  of  the  Indian  and  tropical  western  and  central 
Pacific  Oceans"  (Peters). 

Recorded  in  the  Hawaiian  Group  from  Midway,  Lisianski  and  Laysan 
(breeding),  French  Frigate  Shoal  (Bryan),  "Bird  Island"  (Nihoa) 
(breeding),  Lehua  (Rothschild),  Niihau,  but  see  W.  K.  Fisher,  Bull. 
U.  S.  Fish  Comm.,  23, 1903,  p.  797,  Kaula,  Kauai,  Oahu  (on  Mokumanu 
Island)  (breeding),  and  many  sight  records  at  sea. 

Sula  leucogaster  plotus  (Forster) 

Pelecanus  plotus  Forster,  Descriptiones  Animalium,  etc.,  (ed.  Lichtenstein) 

1844,  p.  278.    (Near  New  Caledonia.) 
Booby,  Common  Brown  Booby,  Brown  Vested  Booby 
Sula  sula  Linn,  in  Rothschild,  Avifauna  of  Laysan,  etc.,  pp.  28,  297;  Wilson 

and  Evans,  Aves  Hawaiiensis,  p.  xxv;  Henshaw,  Birds  Hawaiian  Islands, 

p.  114;  W.  A.  Bryan,  Key,  Birds  Hawaiian  Islands,  p.  15;  Henshaw, 

Birds  Hawaiian  Possessions,  p.  114. 

Range.  "Western  and  central  tropical  Pacific  to  northeastern 
Australia"  (Peters). 

Recorded  in  the  Hawaiian  Group  from  Midway,  Lisianski  and  Laysan 
Islands,  French  Frigate  Shoal  (sight)  (Fisher),  Necker  (breeding), 
"Bird  Island  "(Nihoa)  (breeding),  Kaula  (Caum),  and  many  sight 
records  at  sea  off  Niihau,  Oahu  (Mokumanu  Islet)  (Munro). 

Sula  dactylatra  person  at  a  Gould 

Sida  personata  Gould,  Proc.  Zool.  Soc.  London,  1846,  p.  21.  (North  and 
northeast  coasts  of  Australia  =  Raine  Island,  North  Queensland,  fide 
Mathews.) 

Booby,  Blue  Faced  Booby,  Masked  Booby,  Masked  Gannet 

Sula  cyanops  Sundevall  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  25;  Wilson 
and  Evans,  Aves  Hawaiiensis,  p.  xxv;  Henshaw,  Birds  Hawaiian  Islands, 
p.  113;  W.  A.  Bryan,  Key,  Birds  Hawaiian  Group,  p.  15. 

Range.  "Breeds  on  islands  of  the  central  and  western  tropical 
Pacific  Ocean"  (Peters). 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Lisianski, 
Laysan  (breeding),  French  Frigate  Shoal  (breeding),  Necker  (breed- 
ing), "Bird  Island"  (Nihoa)  (breeding),  Kaula,  Lehua  (Caum). 

Family  PHALACROCORACIDAE 

Genus  Phalacrocorax — Cormorants 

Phalacrocorax  Brisson,  Ornithologie,  1,  1760,  p.  60.  Type,  by  tautonomy, 
Phalacrocorax  =  Pelecanus  carbo  Linnaeus. 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  99 

Phalacrocorax  pelagicus  pelagic  is  Pallas 

Phalacrocorax  pelagicus  Pallas,  Zoographia  Rosso-Asiatica,  2,   1811,  p.  303. 

(Eastern  Kamchatka  and  the  Aleutian  Islands.) 
Pelegic  Cormorant 

Range.  Breeds  on  the  coasts  of  northeastern  Asia,  Aleutian  Islands 
to  south-central  Alaska;  south  in  winter  to  Japan  and  Puget  Sound. 

Recorded  in  the  Hawaiian  Group  {as  an  accidental  visitor)  from 
Laysan  1896  (Rothschild),  ?Hilo,  Hawaii  1900  (recorded  by  Henshaw 
with  doubt). 

Family  FREGATIDAE 

Genus  Fregata — Frigate  Birds  or  Man  o'War  Birds 

Fregata  minor  palmerstoni  (Gmelin) 

Pelecanns  palmerstoni  Gmelin,   Systema    Naturae,   1,  pt.   2,    1789,   p.   573. 

(Palmerston  Island  =  Palmerston  Atoll,  Cook  Group.) 
Man  o'War  Bird,  Ima 
Fregata  aquila  Linn,  in  Rothschild,  Avifauna  of  Laysan,  etc.,  pp.  22,  297  and 

other  works. 

Range.     "Hawaiian  Islands  south  to  New  Zealand"  (Peters). 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Pearl  and 
Hermes  Reef,  Laysan  (breeding),  French  Frigate  Shoal,  Necker 
(breeding),  "Bird  Island"  (Nihoa)  (breeding),  Kaula,  Lehua,  Kauai, 
Oahu. 

Order  CICONIIFORMES 

Family  ARDEIDAE— Herons 

Genus  Nycticorax — Night  Herons 

Nycticorax  T.  Forster,  Synoptic  Catalog  British  Birds,  1817,  p.  59.  Type,  by 
tautonomy  and  monotypy,  Nycticorax  infaustus  Forster  =  Ardea  nycti- 
corax Linn. 

Nycticorax  nycticorax  hoactli  (Gmelin) 

Ardea  hoactli  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  630.    (In  Novae 

Hispania  lacubus  =  Valley  of  Mexico.) 
Black  Crowned  Night  Heron,  Auku  Aukuu,  Auku  Kahili  or  Auku  Kahili 
Nycticorax  nycticorax  naevius  (Boddaert)  in  Rothschild,  Avifauna  of  Laysan, 

etc.,  p.  265. 
Nycticorax  griseits  (Boddaert)  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  201, 

and  Henshaw,  Birds  Hawaiian  Islands,  p.  102. 


100  bulletin:  museum  of  comparative  zoology 

Range.  Northern  United  States  and  northeastern  Canada  south 
through  Mexico  and  Central  America  and  east  coast  of  South  America 
to  eastern  Argentine. 

Recorded  in  the  Hawaiian  Group  from  Midway  (skin  in  Bishop 
Museum — D.  R.  Chisholm  coll.),  Oahu,  Molokai,  Hawaii — "all 
islands"  (Rothschild,  AYilson  and  Evans,  Henshaw). 

The  records  of  Demigretta  sacra  seem  to  be  doubtful.  As  Bryan 
(1901,  p.  21)  remarks,  Dole's  record  of  1879  might  just  as  well  refer 
to  the  Auku. 

The  identity  of  the  White  Heron  which  Dr.  Finsch  saw  so  long  ago 
at  Kahalui  is  problematical. 


Family  THRESKIORNITHIDAE 

Genus  Plegadis — Glossy  Ibises 

Plegadis  Kaup,  Skizzirte  Entwickelungs-Geschichte  Natiirliches  System,  etc., 
1829,  p.  82.   Type,  by  monotypy,  Tantalus  falcinellus  Linnaeus. 

Plegadis  guarauna  (Linnaeus) 

Scolovax  gaurauna  Linnaeus,   Systema  Naturae,   ed.    12,   1,    1766,   p.   242. 

(Brazil.) 
White  Faced  Glossy  Ibis 

Range.     Northwestern  North  America  south  to  Argentina. 
Recorded  in  the  Hawaiian  Group   (as  an  accidental  visitor)   from 
Kauai,  ?Maui  (Henshaw),  Molokai  (Munro). 


Order  ANSERIFORMES 

Family  ANATIDAE 

Genus  Chen — Snow  Geese 

Chen  Boie,  Isis  von  Oken,  10,  1822,  column  563.   Type,  by  monotypy,  Anser 
hyperboreus  Pallas. 

Chen  hyperborea  hyperborea  (Pallas) 

Anser  hyperboreus  Pallas,  Spicilegia  Zoologica,  etc.,  1767  (1769?),  fascicule  6 

1769,  p.  31.    (Northeastern  Siberia.) 
Lesser  Snow  Goose 


BRYAN  AM)  GKKK.WVAY:  HAWAIIAN    BIRDS  101 

Range.  Arctic  North  America  south  in  winter  to  temperate  zone. 
Probably  also  Siberia  south  in  winter  to  Japan. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Maui  by  Rothschild  who  received  a  specimen  collected  there  by 
Brother  Matthias  Newell;  Oahu  (J.  E.  Whitney  coll.). 

Genus  Axser — Geese 

A  user  Brisson,  Ornithologie,   1,   1760,  p.   58.    Type,   by  tautonomy,   Anser 
domesticus  =  Anser  anser  Linnaeus. 

Anser  albifrons  gambelli  Hartlaub 

Anser  gambelli  Hartlaub,  Revue  et  Magasin  Zoologie,  etc.,  1852,  p.  7.    (Texas 

and  southern  United  States.) 
American  White-Fronted  Goose 

Range.  Northwestern  North  America  south  in  winter  to  northern 
California. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
"a  lake  on  Mr.  Clark's  estate  at  Honokaohau,"  Hawaii,  where  it  was 
shot  December  18,  1891  (Rothschild),  Molokai  (G.  C.  Munro  coll.). 

Genus  Philacte — Emperor  Goose 

Philacte  Bannister,  Proceedings  Acad.  Nat.  Sci.  Philadelphia,  1870,  p.  131. 
Type,  by  monotypy,  Anas  canagica  Sewastianov. 

Philacte  canagica  (Sewastianov) 

Anas  canagica  Sewastianov,  Nova  Acta  Acad.  Sci.  Imp.  St.  Petersburg,  13, 

1802,  p.  349,  pi.  10.    (Kanaga  Island,  Aleutian  Islands.) 
Emperor  Goose 

Range.  Breeds  on  coasts  of  Siberia  and  Alaska,  winters  chiefly 
in  the  Aleutian  Islands. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Puna,  Hawaii  (Henshaw  1903). 

Genus  Branta — Brant  or  Brent  Geese 

Branta  Scopoli,  Annus  I,  Historico-Naturalis,  1769,  p.  67.  Type,  by  sub- 
sequent designation,  Anas  bernicla  Linnaeus  (Bannister,  Proceedings 
Acad.  Nat.  Sci.,  Philadelphia,  1870,  p.  131). 


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Branta  bernicla  nigricans  (Lawrence) 

Anser  nigricans  Lawrence,  Annals  Lyceum  Nat.  Hist.  New  York,  4,  1846, 

p.  171,  pi.  12.    (Egg  Harbor,  New  Jersey.) 
Black  Brant 

Range.  Arctic  coasts  of  Siberia  and  Arctic  North  America;  south 
in  winter  to  Japan  and  North  China,  and  to  Lower  California. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Maui 
(Rothschild). 

Branta  canadensis  minima  Ridgway 

Branta  minima  Ridgway,  Proceedings  U.  S.  Nat.  Museum,  8,  1885,  p.  22. 

("Pacific  coasts  of  North  America";  type  from  St.  Michael's,  Alaska.) 
Cackling  Goose 
Bernicla  Munroii  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10,  1892, 

p.  108.   (Kauai.) 

Range.  Breeds  on  the  Bering  Sea  coast  of  Alaska  and  the  Aleutian 
Islands;  in  winter,  west  of  the  Rocky  Mountains  from  southern 
British  Columbia  to  southern  California. 

Recorded  in  the  Hawaiian  Group  (as  an  occasional  or  chance  visitor) 
from  Kauai  (Rothschild),  ?Hawaii  (Henshaw,  q.  v.),  Molokai  (G.  C. 
Munro  coll.). 

Genus  Nesochen — Hawaiian  Goose 

Nesochen  Salvadori,  Catalogue  Birds  British  Mus.,  27,  1895,  pp.  81,  126. 
Type,  by  original  designation  and  monotypy,  Anser  sandvicensis  Vigors. 

Nesocehn  sandvicensis  (Vigors) 

Anser  sandvicensis  Vigors,  List  Animals  Garden  Zool.  Soc,  ed.  3,  1833,  p.  4. 
(Hawaiian  Islands.) 

Hawaiian  Goose,  Nene 

Bernicla  sandvicensis  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  187;  Hen- 
shaw, Birds  Hawaiian  Islands,  p.  103. 

Range.     Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Hawaii.  (Reports  from 
Maui,  Kauai  and  Niihau  have  never  been  confirmed.)  Breeds  on  the 
Kona  coast,  and  above  5000  feet;  from  the  Kona  District  to  the  north- 
east side  of  Mauna  Kea,  descending  to  1000  feet  and  sea  level  in 
winter.  Nests  in  lava  flows  or  open  fields.  Now  much  reduced  in 
numbers,  though  still  breeding  in  a  wild  state  as  well  as  in  captivity. 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  103 


Genus  Anas 

Anas  Linnaeus,  Systema  Naturae,  ed.  10, 1,  1758,  p.  122.  Type,  by  subsequent 
designation,  Anas  boschas  Linnaeus  =  Anas  platyrhynchos  Linnaeus. 


Anas  platyrhynchos  platyrhynchos  Linnaeus 

Anas  platyrhynchos  Linnaeus,  Systema  Naturae,  ed.    10,   1,   1758,  p.    125. 

(Europe,  type  locality  restricted  to  Sweden.) 
Mallard 

Range.     Temperate  zones  of  Europe,  Asia  and  North  America. 
Recorded  in  the  Hawaiian.  Group   (as  an  accidental  visitor)   from 
Laysan  (Schauinsland),  Oahu,  Molokai  (Perkins). 


Anas  wyvilliana  wyvilliana  Sclater 

Anas  wyvilliana  Sclater,  Proc.  Zool.  Soc.  London,  1878,  p.  350.    (Hawaiian 

Islands.) 
Hawaiian  Duck,  Koloa  Maoli 
Anas  aberti  Ridgway,  Proc.  U.  S.  Nat.  Mus.,  1,  1878,  p.  250.    (Mazatlan, 

Mexico.) 

Range.  Hawaiian  Islands.  Recorded  from  Niihau,  Kauai,  Oahu, 
Molokai,  Maui  and  Hawaii  in  ponds  and  lakes  along  the  coast  and 
in  mountain  streams  and  bogs  to  8000  feet. 

Recorded  recently  from  Kauai,  Mokulua  Islets,  N.  E.  coast  of 
Oahu  (breeding)  (Munro),  Molokai,  highlands  of  Hawaii  (Baldwin). 

The  type  of  Anas  aberti  Ridgway  was  probably  an  escaped  captive. 
Mazatlan  was  formerly  a  port  for  vessels  from  China  and  possessed  a 
large  bird  market. 


Anas  wyvilliana  laysanensis  Rothschild 

Anas  laysanensis  Rothschild,  Bull.  British  Ornith.  Club,  1,   1892,  p.  xvii. 

(Laysan  Island.) 
Laysan  Teal 

Range.  Laysan.  Recorded  by  Kittlitz  (Museum  Senckenbergianum 
1834,  p.  124)  from  Lisianski  but  never  confirmed. 

Twenty  birds  left  in  1923  (Wetmore,  Nat'l  Geographic  Mag.,  48, 
no.  1,  1925,  p.  103).  Nine  left  in  1936  (W.  F.  Coultas,  in  Hit). 


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Anas  crecca  carolinensis  Gmelin 

Anas  carolinensis  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  533.  (Carolina  to 

Hudson's  Bay.) 
Green  Winged  Teal 

Range.  Northern  North  America;  in  winter  south  to  West  Indies 
and  Panama. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Laysan  (Schauinsland),  Maui  (G.  P.  Wilder  coll.),  Oahu  (Northwood), 
Molokai  (G.  C.  Munro  coll.). 


Anas  acuta  acuta  Linnaeus 

Anas  acuta  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  126.    (Europe — 

type  locality  restricted  to  Sweden.) 
Pintail,  Koloa  Mapu 

Range.  Europe,  Northwestern  North  America  east  to  Iowa;  south 
in  winter  to  West  Indies,  Panama  and  the  Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  migrant  in  winter) 
from  Laysan,  Kauai,  Oahu  (Northwood),  Hawaii.  "Most  of  the 
islands;  common"  (Perkins).  "Well  known  to  sportsmen  as  a  winter 
visitor"  (Henshaw). 


Genus  Mareca — W7idgeons 

Mareca  Stephens,  in  Shaw,  General  Zoology,  12,  pt.  2,  1824,  p.  30.  Type,  by 
subsequent  designation,  Mareca  fistularis  Stephens  =  Anas  penelope 
Linnaeus  (Eyton,  Monograph  Anatidae,  1838,  p.  33). 


Mareca  Americana  (Gmelin) 

Anas  americana  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  526.    (Louisiana 

and  New  York.) 
American  Widgeon,  Baldpate 

Range.  Northwestern  to  north-central  North  America;  south  in 
winter  to  the  Atlantic  coast,  West  Indies  and  Central  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor  in  winter) 
from  Laysan  (Schauinsland),  Oahu  (Northwood),  Maui  (Perkins), 
Molokai  (Munro  coll.). 


BRYAN  AND  GRKKN WAY :  HAWAIIAN'  KIRDS  105 


Genus  Chaulelasmus — Gadwall  Ducks 

Chaulelasmus  Bonaparte,  Geographical  and  Comparative  List  of  Birds  of 
Europe  and  North  America,  1838,  p.  56.  Type,  by  monotypy,  Anas 
strepera  Linnaeus. 

Chaulelasmus  streperus  (Linnaeus) 

Anas  strepera  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  125.    (Europe — 

type  locality  restricted  to  Sweden.) 
Gad  wall 

Range.     Worldwide  within  the  northern  hemisphere. 
Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor  in  winter) 
from  Oahu  (Perkins),  Molokai  (Munro  coll.). 

Genus  Spatula — Shoveller  Ducks 

Spatula  Boie,  Isis  von  Oken,  1822,  col.  564.  Type,  by  monotypy,  Anas 
clypeata  Linnaeus. 

Spatula  clypeata  (Linnaeus) 

Anas  clypeata  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  124.    (Coasts  of 

Europe;  type  locality  restricted  to  southern  Sweden.) 
Shoveller,  North  West  Duck,  Koloa  Moha 

Range.     Worldwide  within  the  northern  hemisphere. 
Recorded  in  the  Hawaiian  Group  (as  a  regular  migrant  in  winter) 
from  Laysan,  Kauai,  Oahu,  Molokai,  Lanai,  Maui,  Hawaii. 

Genus  Bucephala — Golden  Eyes,  Buffle  Head 

Bucephala  Baird,  Reports,  Explorations  and  Surveys  for  a  Railroad  from  the 
Mississippi  River  to  the  Pacific  Ocean,  9,  pt.  2,  pp.  L,  788,  785.  Type,  by 
original  designation,  Anas  albeola  Linnaeus. 

Bucephala  albeola  (Linnaeus) 

Anas  albeola  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  124.    (America  = 

Newfoundland  ex  Edwards.) 
Bvffle  Head 

Range.  Northwestern  North  America;  in  winter  eastward  and 
southward  in  the  United  States. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Laysan  (Rothschild),  Oahu  (Northwood),  Maui  (Perkins). 


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Genus  Nyroca — Diving  Ducks 

Nyroca  Fleming,  Philosophy  of  Zool.  etc.,  2,  1822,  p.  260.  Type,  by  tautonomy, 
Anas  nyroca  Giildenstadt. 


Nyroca  affinis  (Eyton) 

Fuligula  affinis  Eyton,  Monograph  Anatidae,  1838,  p.  157.    (North  America.) 
Lesser  Scaup  Duck 

Range.  Northern  North  America;  south  in  winter  to  Central 
America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Lanai  (G.  C.  Munro  coll.). 

Genus  Histrionicus — Harlequin  Ducks 

Histrionicus  Lesson,  Manuel  d'Ornithologie,  2,  1828,  p.  415.  Type,  by  original 
designation,  Anas  histrionica  Linnaeus. 

Histrionicus  histrionicus  pacificus  W.  S.  Brooks 

Histrionicus  histrionicus  pacificus  W.   S.   Brooks,   Bull.    Mus.  Comp.  Zool., 

Cambridge,  Mass.,  59,  1915,  p.  393.    (Cape  Shipunski,  Kamchatka.) 
Western  Harlequin  Duck 

Range.  Eastern  Siberia  and  northwestern  North  America ;  south  in 
winter  to  Japan  and  California. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Lay- 
san  (P.  E.  H.  Bompke  coll.  1906). 

Genus  Mergus — Mergansers 

Mergus  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  129.   Type,  by  subse- 
quent designation,  Mergus  castor  Linnaeus  =  Mergus  serrator  Linnaeus. 

Mergus  serrator  Linnaeus 

Mergus  serrator  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  129.    (Europe. 
Type  locality  restricted  to  Sweden.) 

Range.  Worldwide  in  the  northern  hemisphere ;  south  in  winter  to 
temperate  zone. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  0§hu 
(Bryan),  Hawaii  (Henshaw). 


BRYAX  AND  GRKK.WVAY:  HAWAIIAN  BIRDS  107 

Order  FALCOXIFORMES 
Family  ACCIPITRIDAE 

Genus  Buteo — Soaring  Hawks  and  Buzzards 

Buteo  Lacepede,  Discours  .  .  .  Tableaux  methodiques  mammiferes  .  .  • 
Oiseaux,  1799,  p.  4.  Type,  by  tautonom}',  Falco  buteo  Linnaeus. 

Buteo  solitarius  Peale 

Buteo  solitarius  Peale,  U.  S.  Exploring  Expedition,  8,  1848,  p.  62.    (Island  of 

Hawaii.) 
Hawaiian  Hawk,  Io 
Accipiter  hawaii  Dole,  Hawaiian  Almanac  for  1879  (1878)  p.  43. 

Range.     Island  of  Hawaii  (all  parts). 

Genus  Circus — Marsh  Hawks 

Circus  Lacepede.  Discours  .  .  .  Tableaux  methodiques  mammiferes  oiseaux, 
1799,  p.  4.  Type,  by  subsequent  designation,  Falco  aeruginosus  Linnaeus 
(Lesson,  Manuel  d'Ornith.,  1,  1828,  p.  105). 

Circus  cyaneus  hudsonius  (Linnaeus) 

Falco  hudsonius  Linnaeus,  Systema  Naturae,  ed.  12,  1,  1766,  p.  128.  (Hudson 
Bay  ex  Edwards.) 

Marsh  Hawk- 
Range.     Northern  and  central  North  America;  south  in  winter  to 

Central  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Oahu 

(Wilson). 

Genus  Pandion — Ospreys 

Pandion  Savigny,  Description  d'Egypt,  Oiseaux,  1809,  p.  69.  Type,  by 
monotypy,  Pandion  fluvialis  Savigny  =  Falco  haliaetus  Linnaeus. 

Pandion  haliaetus  carolinensis  (Gmelin) 

Falco  carolinensis  Gmelin,  Systema  Naturae,  1,  pt.  1,  1788,  p.  263.    (No  type 

locality.  Carolina  ex  Catesby  et  al.) 
American  Osprey 


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Range.  Northern  North  America;  south  in  winter  to  the  West 
Indies  and  South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  winter  visitor) 
from  ?Niihau  (Dole),  Oahu  (Perkins),  ?Molokai  (Dole),  ?Hawaii 
(Dole). 

Family  RALLIDAE— Rails 

Genus  Porzanula 

Porzanula  Frohawk,  Annals  and  Magazine  Nat.  Hist.,  London,  (6),  9,  1892, 
p.  247.   Type,  by  monotypy,  Porzanula  palmeri  Frohawk. 

Porzanula  palmeri  Frohawk 

Porzanula  Palmeri  Frohawk,  Annals  and  Magazine  Nat.  Hist.,  London  (6),  9, 

1892,  p.  247.    (Laysan  Island.) 
Laysan  Rail,  Laysan  Crake 

Range.  Laysan  Island,  where  but  two  remained  in  1923  (Wetmore 
1925);  Midway  Island,  where  introduced  in  the  latter  part  of  the  19th 
century  (Rothschild,  p.  xiii)  and  where  common  (Denig  et  al). 

Genus  Pennula 

Pennula  Dole,  Hawaiian  Almanac  and  Annual  for  1879  (1878),  p.  54.  Type, 
by  monotypy,  Pennula  millei  (sic)  Dole. 

Pennula  millsi  Dole 

Pennula  millei  [sic]  Dole,  Hawaiian  Almanac  and  Annual  for  1879  (1878), 
p.  54.    (Hawaii.) 

Hawaiian  Rail,  Moho 

Rallus  ecaudotus  (sic)  King,  in  Cook's  Voyage  to  the  Pacific  Ocean,  3,  1784, 
p.  119.  (Hawaiian  Islands);  Pennula  ecaudata  (King)  in  Wilson  and 
Evans,  Aves  Hawaiiensis,  p.  171,  and  in  Perkins,  Fauna  Hawaiiensis,  1, 
pt.  4,  p.  453,  and  older  works.  This  name  is  preoccupied  by  Rallus  ecau- 
datus  Miller  1783. 

Range.  Formerly  Island  of  Hawaii  in  open  grassy  country  or  low 
scrub  just  below  the  heavy  rain  forest,  possibly  in  the  Olaa  district, 
certainly  on  the  windward  side  of  Kilauea  and  perhaps  about  forty 
miles  north  along  the  coast.  ?Molokai  (Perkins).  Extinct,  last  speci- 
men seen  1884  (Rothschild),  ?1893  (Henshaw).  Five  specimens  in 
museums. 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  109 

Pennula  sandwichensis  (Gmelin) 

Rallus  sandwichensis  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  717.  ("in 
insulis  Sandwich,"  based  on  Latham's  Sandwich  Rail.) 

Spotted  Hawaiian  Rail 

Rallus  obscurus  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  718.  ("in  insulis 
Sandwich,"  based  on  Latham's  Dusky  Rail,  and  other  authors.) 

Pennula  wilsoni  Finsch,  Notes  Leyden  Mus.,  20,  1898,  p.  77.  Hawaiian 
Islands.) 

Range.  Unknown.  Now  extinct.  Known  only  from  the  unique 
type  in  Leyden,  Holland.  Paler  than  millsi  with  central  black  spot  on 
feathers  of  back. 

Genus  Gallinula — Coots,  Gallinules 

Gallinula  Brisson,  Ornithologie,  1,  1760,  p.  2.  Type,  by  tautonomy,  Gallinula 
Brisson  =  Fulica  chloropus  Linnaeus. 

Gallinula  chloropus  sandvicensis  Streets 

Gallinula  sandvicensis  Streets,  Ibis,  1877,  p.  25.    (Oahu,  Hawaiian  Islands.) 
Hawaiian  Gallinule,  Alae,  Alae  Ula 

Range.  Hawaiian  Islands.  Recorded  from  Kauai,  Oahu,  Molokai, 
Maui,  Hawaii  in  ponds,  rice  fields  and  taro  patches. 

Genus  Fulica — Coots 

Fulica  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  152.  Type,  by  tau- 
tonomy, Fulica  atra  Linnaeus. 

Fulica  Americana  alai  Peale 

Fulica  alai  Peale,   United  States  Exploring  Expedition,   8,    1848,   p.   224. 

(Hawaiian  Islands.) 
Hawaiian  Coot,  Alae  Keokeo 

Range.  Hawaiian  Islands.  Recorded  from  Niihau,  Kauai,  Oahu> 
Molokai,  Maui,  Hawaii. 


Order  CHARADRIIFORMES 

Family  CHARADRIIDAE 

Genus  Squatarola — Plovers 

Squatarola  Cuvier,  Regne  Animal,  1,  1817  (1816),  p.  467.  Type,  by  tautonomy, 
Tringa  squatarola  Linnaeus. 


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Squatarola  squatarola  (Linnaeus) 

Tringa  squatarola  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  149.   (Europe. 

Type  locality  restricted  to  Sweden  by  Hartert,  Vogel  palaarktischen 

Fauna,  2,  p.  1553.) 
Black-bellied  Plover,  Beetle  Head 

Range.  Circumpolar  in  the  arctic;  south  in  winter  to  South  Africa* 
India,  Australia,  western  South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor  on  migration) 
from  Oahu  (Northwood  et  al),  Hawaii  (Henshaw). 

Genus  Pluvialis 

Pluvialis  Brisson,  Ornitholigie,  1760,  1,  p.  46;  5,  p.  42.  Type,  by  tautonomy, 
Pluvialis  aurea  Brisson  =  Charadrius  pluvialis  Linnaeus  =  Charadrius 
apricarius  Linnaeus. 

Pluvialis  dominica  fulva  (Gmelin) 

Charadrius  fulvus  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  687.    (Tahiti.) 

Pacific  Golden  Plover,  Kolea 

Charadrius  fulvus  in  Rothschild  and  other  works. 

Range.  Siberia  and  Alaska;  south  in  winter  to  India,  Australia, 
Indo-China,  Pacific  Islands. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  winter  visitor)  from 
Midway,  Laysan,  Lisianski,  French  Frigate  Shoal,  Bird  Island  (Nihoa), 
Niihau,  Kauai,  Oahu,  Molokai,  Maui,  Hawaii,  "all  islands"  (Perkins). 
Arrive  in  August  or  September;  depart  in  April  or  May  as  a  rule,  but 
sometimes  they  remain  during  the  summer,  though  probably  they 
never  breed. 

Genus  Charadrius — Plovers 

Charadrius  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  150.  Type,  by 
tautonomy,  Charadrius  hiaticula  Linnaeus. 

Charadrius  vociferus  vociferus  Linnaeus 

Charadrius  vociferus  Linnaeus,  Systema  Naturae,  ed.   10,  1,   1758,  p.   150. 

(North  America  =  South  Carolina  ex  Catesby.) 
Killdeer 

Range.  Arctic  and  temperate  zones  and  portions  of  the  eastern 
tropical;  south  in  winter  to  northern  South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Maui 
(W.  A  .Bryan). 


BRYAN    AND  GREENWAY:   HAWAIIAN    BIRDS  111 

Family  SCOLOPACIDAE 

Genus  Numenius — Curlews 

Numenius  Brisson,  Ornithologie,  17G0, 1,  p.  48;  5,  p.  311.  Type,  by  tautonomy, 
X miliums  Brisson   =  Scolopax  arquata  Linnaeus. 

Numenius  tahitiensis  (Gmelin) 

Scolopax  tahitiensis  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  656.    (Tahiti 

ex  Latham.) 
Bristle  Thighed  Curlew,  Kioea 

Range.     Alaska;  south  in  winter  to  the  Pacific  islands. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  winter  visitor)  from 
.Midway,  Laysan,  Niihau,  Kauai,  Oahu,  Molokai,  Lanai  (sight) 
(Rothschild),  Maui,  Hawaii.  Like  the  Golden  Plover  (Kolea),  this 
bird  leaves  in  April  or  May  and  arrives  in  August  or  September,  but 
sometimes  remains  during  the  summer,  though  it  probably  never 
breeds  on  the  islands. 

Genus  Limosa — Godwits 

Limosa  Brisson,  Ornithologie,  1760,  1,  p.  48;  5,  p.  261.  Type,  by  tautonomy, 
Limosa  Brisson  =  Scolopax  limosa  Linnaeus. 

Limosa  lapponica  baueri  Naumann 

Limosa  baueri  Naumann,  Naturgeschichte  Vogel  Deutschlands,  8,  1836,  p. 
429.    (New  Holland.) 

Pacific  Godwit 

Limosa  lapponica  novaezealandiae  Gray  in  Rothschild,  Avifauna  of  Laysan, 
etc.,  p.  307;  Henshaw,  Birds  Hawaiian  Islands,  etc.,  p.  93;  Bryan,  Key 
Birds  Hawaiian  Group,  p.  27;  Perkins,  Fauna  Hawaiiensis  (Aves),  p.  45. 

Range.  Northeastern  Asia  and  northwestern  North  America; 
south  in  winter  to  islands  of  the  Pacific  and  Australia. 

Recorded  in  the  Hawaiian  Group  (as  an  occasional  visitor  on  migration) 
from  Laysan  (Schauinsland),  Kauai  (W.  A.  Bryan),  Maui  (Ridgway). 

Genus  Heteroscelus — Tatlers 

Heteroscelus  Baird,  Reports,  Explorations  and  Surveys  for  a  Railroad  from  the 
Mississippi  River  to  the  Pacific  Ocean,  9,  1858,  pp.  xxii,  xlvii,  728,  734. 
Type,  by  monotypy,  Totanus  brevipies  Vieillot. 


112  bulletin:  museum  of  comparative  zoology 

Heteroscelus  incanus  (Gmelin) 

Scolopax  incana  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  658.    (Eimeo 

(=  Moorea)  and  Palmerston  Islands.) 
Wandering  Tattler,  Ulili 
Heteractitis  incanus  (Gmelin)  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  255; 

Henshaw,  Birds  Hawaiian  Islands,  etc.,  p.  92;  Bryan,  Key  Birds  Hawaiian 

Group,  p.  27;  Perkins,  Fauna  Hawaiiensis  (Aves),  p.  450;  Totanus  incanus 

in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  151. 

Range.  Alaska;  south  in  winter  to  the  west  coast  of  America  to 
Ecuador  and  to  the  islands  of  the  Pacific,  sometimes  to  Australia. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  winter  resident)  from 
Midway,  Laysan,  Lisianski,  Necker,  French  Frigate  Shoal,  Niihau, 
Kauai,  Oahu,  Maui,  Hawaii,"  ....  frequenting  the  rocky  shores  of  all 
islands"  (Henshaw).  Usually  arrive  in  August  and  leave  in  May. 
Though  certain  birds  may  remain  all  summer,  there  is  no  breeding 
record  for  the  islands. 

Genus  Arenaria — Turnstones 

Arenaria  Brisson,  Omithologie,  1760,  1,  p.  48;  5,  p.  132.  Type,  by  tautonomy, 
Arenaria  Brisson  =  Tringa  interpret  Linnaeus. 

Arenaria  interpres  interpres  (Linnaeus) 

Tringa  interpres  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  149.  (Europe 
and  North  America;  type  locality  restricted  to  Gotland  Island,  Sweden, 
by  Hartert,  Vogel  Palaartischen  Fauna,  p.  1566.) 

Turnstone,  Akekeke 

Tringa  oahuensis  Bloxam,  Voyage  "Blonde",  1826,  p.  132. 

Range.  Northern  Europe  and  northern  Asia  and  Alaska  west  of 
Point  Barrow;  south  in  winter  to  Africa,  India,  China  and  islands  of 
the  Pacific. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  winter  resident)  from 
Midway,  Lisianski,  Laysan,  French  Frigate  Shoal,  Necker,  Niihau, 
Kauai,  Oahu,  Molokai,  Maui,  Hawaii,  "All  the  islands"  (Perkins). 
Usually  arrive  in  August  or  September  and  depart  in  April  or  May. 
Certain  birds  remain  all  summer,  some  in  flocks  at  6000  feet  on  Halea- 
kala,  Maui  and  in  the  crater  of  Kilauea,  Hawaii.  Subspecific  identifica- 
tion of  winter  birds  is  impossible.  It  seems  more  probable  that  most 
of  the  birds  to  be  seen  in  the  Hawaiian  Islands  are  interpres,  though  in- 
dividuals of  morinella,  which  is  the  bird  that  breeds  in  North  America 
east  of  Point  Barrow,  Alaska,  may  also  reach  the  islands. 


BRYAN  AND  GREENWAY:  HAWAIIAN'  BIRDS  113 

Genus  Capella — Snipe 

Capella  Frenzel,  Beschreibung  Vogel  und  Eyer  .  .  .  Wittenborg  .  .  .  ,  1801, 
p.  58.  Type,  by  nionotypy,  Scolopax  coelestis  Frenzel  =  Scolopax  gallinago 
Linnaeus. 

(  apella  delicata  (Ord) 

Scolopax  delicata  Ord  in   Wilson,   American   Ornithology,   9,   1825,   p.   218. 

(Pennsylvania.) 
Wilson's  Snipe,  Jack  Snipe 
Gallinago  delicata  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  253;  Henshaw, 

Birds  Hawaiian  Islands,  p.  94;  Perkins,  Fauna  Hawaiiensis  (Aves),  p.  451. 

Range.  Arctic  and  temperate  North  America;  south  in  winter  to 
South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  occasional  visitor  on  migration) 
from  Laysan,  Oahu,  Molokai,  Maui,  Hawaii. 


Genus  Crocethia — Sanderling 

Crocethia  Billberg,  Synopsis  Fauna  Scandanaviae,  1,  pt.  2,  1828,  p.  132.  Type 
by  nionotypy,  Charadrius  calidris  Linnaeus  =  Trynga  alba  Pallas. 


Crocethia  alba  (Pallas) 

Trynga  alba  Pallas  in  Vroeg,  Beredeneerde  Catalogus,  etc.,  Adumbratiunculae, 
1764,  N.  320,  p.  7.  ("Noordsche  Zeekusten"  =  coast  of  northern  Holland.) 

Sanderling,  Hunakai 

Calidris  aretiaria  (Linnaeus)  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  259; 
Wilson  and  Evans,  Aves  Hawaiiensis,  p.  153,  159;  Henshaw,  Birds 
Hawaiian  Islands,  etc.,  p.  93;  Perkins,  Fauna  Hawaiiensis  (Aves),  p.  451, 
and  other  works. 

Range.  Arctic  coasts  of  Europe,  Asia,  America ;  south  in  winter  to 
temperate  zone  and  southern  hemisphere — Cape  of  Good  Hope, 
Malaya,  Australia,  Islands  of  the  Pacific,  southern  South  America. 

Recorded  in  the  Hawaiian  Group  (as  a  regular  winter  resident)  from 
Laysan,  Niihau,  Kauai,  Maui  (Ridgway),  Oahu,  Molokai,  Hawaii, 
"Most,  and  probably  all,  islands"  (Perkins). 

We  have  followed  the  A.  O.  U.  Check-list  and  Peters  (Birds  of  the 
^Yorld)  in  recognizing  but  one  form  of  this  species.  If  more  are  recog- 
nized the  name  of  the  bird  which  visits  the  islands  will  be  Crocethia 
alba  tridactyla  Pallas,  the  supposed  form  which  breeds  in  northwestern 
North  America. 


114  bulletin:  museum  of  comparative  zoology 

Genus  Erolia — Sandpipers 

Erolia  Vieillot,  Analyse  nouvelle  ornithologie  .  .  .  1816,  p.  55.    Type,  by 
monotypy,  Erolia  variegata  Vieillot  =  Scolopax  testacea  Pallas. 

Erolia  acuminata  (Horsfield) 

Totanus  acuminatum  Horsfield,  Transactions  Linnaean  Soc.  London,  13,  pt.  1, 

1821,  p.  192.    (Java.) 
Sharp-tailed  Sandpiper 
Heteropygia  acuminata  in  Rothschild,  Avifauna  Laysan,  etc.,  p.  255,  307; 

Tringa  acuminata  in  Henshaw,  Birds  Hawaiian  Islands,  p.  94;  Perkins, 

Fauna  Hawaiiensis  (Aves),  p.  451. 

Range.  Eastern  Asia;  south  in  winter  to  south  Pacific  islands, 
Malaya,  New  Guinea,  Australia. 

Recorded  in  the  Hawaiian  Group  (as  an  occasional  visitor  on  migra- 
tion) from  Midway,  Laysan,  Kauai,  Oahu,  Molokai  (specimen  in 
Bishop  Museum),  Maui,  Hawaii. 

Erolia  melanotos  (Vieillot) 

Tringa  melanotos  Vieillot,  Nouveau  Dictionnaire  Hist.  Nat.,  34,  1819,  p.  462. 

(Paraguay.) 
Pectoral  Sandpiper 
Tringa  maculata  in  Henshaw,  Birds  Hawaiian  Islands,  etc.,  p.  94;  Perkins, 

Fauna  Hawaiiensis  (Aves),  p.  451. 

Range.  Arctic  zone  of  eastern  Asia  and  North  America;  south  in 
winter  to  South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor  on  migration) 
from  Oahu  (W.  A.  Bryan),  Hawaii. 

Genus  Himantopus— Stilts 

Himantopus  Brisson,  Ornithologie,  1760,  1,  p.  46;  5,  p.  33.    Type,  by  tau- 
tonomy,  Himantopus  Brisson  =  Charadrius  himantopus  Linnaeus. 

Himantopus  himantopus  knudseni  Stejneger 

Himantopus  knudseni  Stejneger,  Proceedings  U.  S.  Nat.  Mus.,  10,  1887,  p.  81, 

pi.  6,  fig.  2.    (Kauai,  Hawaiian  Islands.) 
Hawaiian  Stilt,  Aeo,  Kukuluaeo 

Range.  Hawaiian  Islands,  where  recorded  from  Niihau,  Kauai, 
Oahu,  Molokai,  Maui  near  lakes  and  ponds  or  on  exposed  mud  flats. 
"I  have  never  seen  it,  however,  upon  Hawaii,  nor  have  I  been  able  to 
learn  of  its  presence  there"  (Henshaw). 


B  RY  A  N  A  ND  GR  E E  N  W  A  V  :  1 1 A  W  AIIAN  BIRDS  11  5 

Family  PHALAROPODIDAE 

Genus  Phalaropus — Phalaropes 

Phalaropus  Brisson,  Ornithologie,  1760,  1,  p.  50;  6,  p.  12.  Type,  by  tautonomy, 
Phalaropus  Brisson  =  Tringa  fulicaria  Linnaeus. 

Phalaropus  fulicarius  (Linnaeus) 

Tringa  fulicaria  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  148.    (Hudson 

Bay  ex  Edwards.) 
Red  Phalarope 
Crymophilus  fulicarius  in  Rothschild,  Avifauna  Laysan,  etc.,  p.  251;  Henshaw, 

Birds  Hawaiian  Islands,  p.  95;  Bryan,  Key  Birds  Hawaiian  Group,  p.  25. 

Range.  Circumpolar  in  arctic;  south  in  winter  to  seas  off  the  coasts 
of  West  Africa  and  Chile. 

Recorded  in  the  Hawaiian  Group  (as  an  irregular  visitor  on  migration) 
from  Laysan,  Kauai,  Oahu,  Maui,  Hawaii,  "evidently  accompanied 
the  Akekeke  (Arenaria  interpres)  to  the  uplands  to  feed"  (!)  (Hen- 
shaw). 

Phalaropus  lobatus  (Linnaeus) 

Tringa  tobata  {Lobata  in  Emendanda,  p.  824)  Linnaeus,  Systema  Naturae,  ed. 

10,  1,  1758,  p.  148.    (Hudson  Bay  ex  Edwards.) 
Northern  Phalarope 

Range.  Circumpolar  in  Arctic  Zone;  south  in  winter  to  oceans  in 
southern  hemisphere. 

Recorded  in  Hawaiian  Group  (as  a  chance  visitor  on  migration)  from 
Kauai. 

Family  LARIDAE 

Genus  Larus — Gulls 

Larus  Linnaeus,  Systema  Naturae,  ed.  10, 1,  1758,  p.  136.  Type,  by  subsequent 
designation,  Larus  marinus  Linnaeus  (Selby,  Catalogue  generic  and  sub- 
generic  types  .  .  .  Aves,  1840,  p.  48). 

Larus  delawarensis  Ord 

Larus  delawarensis  Ord  in  Guthrie's  New  Geographical  .  .  .  Grammar,  etc., 
2nd  American  ed.,  2,  1815,  p.  319.   (Delaware  River,  below  Philadelphia.) 
Ring-billed  Gull 


116  bulletin:  museum  of  comparative  zoology 

Range.  Northwestern  and  northeastern  North  America;  south  in 
winter,  sometimes  as  far  as  the  Greater  Antilles  and  Mexico. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Molokai  and  Maui  or  Hawaii  (Bryan). 

Larus  argentatus  smithsonianus  Coues 

Larus  smithsonianus  Coues,  Proceedings  Academy  Nat.  Sci.,  Philadelphia, 

1862,  p.  296.    (Eastern  and  western  coasts  of  North  America.) 
Herring  Gull 

Range.  North  America  in  arctic  and  northern  temperate  zones; 
south  in  winter,  sometimes  to  northern  tropics. 

Recorded  in  Hawaiian  Group  (as  an  accidental  visitor)  from  Laysan. 

Larus  californicus  Lawrence 

Larus  californicus  Lawrence,  Annals  Lyceum  Nat.  Hist.,  New  York,  6,  1854, 

p.  79.    (Near  Stockton,  California.) 
California  Gidl 

Range.  Northwestern  North  America  from  the  Mackenzie  River 
east  to  Saskatchewan,  south  to  west  central  California  and  north- 
western Wyoming;  south  in  winter  to  southern  California  and  western 
Mexico. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Maui 
or  Hawaii  (Bryan,  1901). 

Larus  pipixcan  Wagler 

Larus  pipixcan  Wagler,  Isis  von  Oken,  1831,  col.  515.    (Mexico.) 
Franklin's  Gull 

Larus  franklini  Swainson  and  Richardson  in  Henshaw,  Birds  of  Hawaiian 
Islands,  p.  127;  Bryan,  Key  Birds  Hawaiian  Group,  p.  6. 

Range.  Interior  of  northwestern  North  America;  south  in  winter 
to  coasts  of  the  Gulf  of  Mexico  and  western  coasts  of  South  America. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Maui 
(one  record). 

Larus  glaucescens  Naumann 

Lams  glaucescens  Naumann,  Naturgeschichte  Vogel  Deutschlands,  etc.,  10, 

1840,  p.  351.    ("Nord-Amerika".) 
Glaucous-uinged  Gull 


BRYAX  AM)  GREENWAT:  HAWAIIAN  BIRDS  117 

Range.  Northeastern  Asia  arid  northwestern  North  America; 
south  in  winter  to  China  and  Japan  and  the  Aleutian  Islands  to  Cali- 
fornia. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Lay- 
san,  Hawaii,  "Probably  .  .  .  other  islands,  especially  to  Oahu"  (Hen- 
shaw). 

This  gull  sometimes  follows  ships  ta  the  Hawaiian  Islands. 

Larus  hyperborevs  Gunnerus 

Larus  hyperboreus  Gunnerus,  in  Leem  .  .  .  Beskrivelse  Finmarkens  Lapper, 

etc.,  1767,  p.  226,  (footnote).    (Northern  Norway.) 
Glaucous  Gull,  Point  Barrow  Gull 

Larus  glaucus  Brunnich,  Ornithologia  borealis,  etc.,  1764,  p.  44. 
Larus  barrorianus  Ridgway,  Auk,  3,  1886,  p.  330;  in  Henshaw,  Birds  Hawaiian 

Islands,  p.  126;  Bryan,  Key  Birds  Hawaiian  Group,  p.  6. 

Range.  Circumpolar  in  the  Arctic;  south  in  winter  to  coast  of 
temperate  zones,  straying  sometimes  far. 

Recorded  in  Hawaiian  Group  (as  an  accidental  visitor)  from  Laysan, 
Kauai,  Lanai,  Maui. 

Larus  Philadelphia  (Ord) 

Sterna  Philadelphia  Ord  in  Guthrie's  New  Geographical  .  .  .  Grammar,  etc., 
2nd  American  ed.,  2,  1815,  p.  319.  (No  locality  =  near  Philadelphia, 
Pennsylvania.) 

Bonaparte  s  Gull 

Range.  Northwestern  North  America ;  winters  on  the  Atlantic  and 
Pacific  coasts. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from  Kauai 
(Rothschild).  ' 

Genus  Rissa — Kitti wakes 

Rissa  Stephens,  in  Shaw's  General  Zoology,  13,  pt.  1,  1826,  p.  180.  Type,  by 
monotypy,  Rissa  brunnichii  Stephens  =  Larus  tridactylus  Linnaeus. 

Rissa  tridactyla  pollicaris  Ridgway 

Rissa  tridactyla  pollicaris  attributed  to  Stejneger  by  Ridgway  in  Baird, 
Brewer  and  Ridgway,  Water  Birds  North  America,  2,  1884,  p.  202. 
(Kotzebue  Sound,  Alaska.) 

Western  Kittiwake 


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Range.  Eastern  Siberia,  Islands  of  Behring  Sea,  Alaska;  south  in 
winter  to  Japan  and  California. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Laysan  (fragments  in  Bishop  Museum). 

Genus  Sterna — Terns 

Sterna  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  137.    Type,  by  tau- 
tonomy,  Sterna  hirundo  Linnaeus. 

Sterna  paradisaea  Pontoppidan 

Sterna  Paradisaea  Pontoppidan,  Danske  Atlas,  1,  1763,  p.  622.    (Christansoe, 

Denmark  ex  Briinnich  1764.) 
Arctic  Tern 

Range.  Circumpolar  in  Arctic  and  parts  of  northern  temperate 
zones;  south  in  winter  to  the  Antarctic  Ocean. 

Recorded  in  the  Hawaiian  Group  (  as  an  accidental  visitor)  from 
Oahu,  Hawaii  (Henshaw). 

Sterna  sumatrana  sumatrana  Raffles 

Sterna  sumatrana  Raffles,  Transactions  Linnaean  Soc.  London,  13,  pt.  2,  1822, 

p.  329.    (Sumatra.) 
Black  naped  Tern 
Sterna  melanauchen  Temminck  in  W.  A.  Bryan,  Birds  Hawaiian  Group,  p.  8; 

Perkins,  Fauna  Hawaiiensis  (Aves),  p.  464;  Henshaw,  Birds  Hawaiian 

Islands,  p.  124. 

Range.     Western  and  southern  Pacific  and  eastern  Indian  Oceans. 
Recorded  in  the  Hawaiian   Group   (as  an  accidental  visitor)  from 
Kauai,  Hawaii. 

Sterna  anaethetus  lunata  Peale 

Sterna  lunata  Peale,  U.  S.  Exploring  Expedition,  8,  1848,  p.  277.    (Vincennes 

Island  =  Kauehi  Island,  Tuamotus.) 
Bridled  Tern,  Gray  backed  Tern,  Pakalakala 
Haliplana  lunata  in  Rothschild,  Avifauna  of  Laysan,  etc.,  v,  vii,  ix,  37. 

Range.  Pacific  Ocean  from  the  Hawaiian  Islands  south  to  the 
Moluccas. 

Recorded  in  the  Hawaiian  Group  from  Laysan  (breeding),  ?  Lisianski, 
Gardner  (sight,  Rothschild),  ?  French  Frigate  Shoal  (Fisher  ?  ex- 
Rothschild),  Necker,  Niihau,  Kauai,  Oahu,  Hawaii. 


BRYAN  AND  GREEXWAY  :  HAWAIIAN  BIRDS  ]  1!) 

Sterna  fuscata  oahuensis  Bloxam 

Sterna  Oahiu  nsis  Bloxam,  Voyage  "Blonde",  1826,  p.  251.   (Oahu.) 

Wideawake,  Sooty  Tern,  Ewa<  wa 

Haliplana  fuMginosa.  (Gmelin)  in  Rothschild,  Avifauna  of  Laysan,  etc.,  vii, 
viii,  39;  Sterna  fuliginosa  (Gmelin)  in  Wilson  andEvans,  AvesHawaiiensis- 
p.  137;  Bryan,  Key  Birds  Hawaiian  Group,  p.  8;  Perkins,  Fauna  Hawai- 
iensis  (Aves),  p.  464;  Henshaw,  Birds  Hawaiian  Islands,  p.  122,  et  al. 

Range.  Pacific  Ocean  from  the  Bonin  Islands  and  the  Hawaiian 
Islands  southward. 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Pearl  and 
Hermes  Reef,  Lisianski,  Laysan  (breeding),  ?  Gardner  (sight,  Roths- 
child), French  Frigate  Shoal  (breeding),  Necker  (breeding),  Bird  Rock 
(Nihoa),  Kaula,  Kauai,  Oahu  (Moku  Manu). 

Genus  Procelsterna — Terns 

Procelsterna  Lafresnaye,  Magazin  Zoologie,  1842,  Oiseaux,  pi.  29,  p.  1.  Type, 
by  monotypy,  Procelsterna  tereticollis  Lafresnaye  =  Sterna  teretirostris 
Lafresnaye. 

Procelsterna  certjlea  saxatilis  W.  K.  Fisher 

Procelsterna  saxatilis  W.  K.  Fisher,  Proceedings  U.  S.  Nat.  Mus.,  26,  1903, 

p.  559.    (Necker  Island.) 
Necker  Island  Tern 

Range.     Marcus  Island  and  western  Hawaiian  Islands. 
Recorded  in  the  Hawaiian  Group  from  French  Frigate  Shoal,  Necker 
(breeding),  Bird  Island  (Nihoa),  Kaula. 

Genus  Ano  us  Stephens — Noddies 

Anoiis  Stephens,  in  Shaw's  General  Zoology,  13,  pt.  1,  1826,  p.  139.  Type,  by 
subsequent  designation,  Anoiis  niger  Stephens  =  Sterna  stolida  Linnaeus 
(Gray,  List  Genera  Birds,  1840,  p.  79). 

Anous  stolidus  pileatus  (Scopoli) 

Sterna  pileata  Scopoli,  Deliciae  florae  et  faunae  insubricae,  etc.,  fascic.  2, 
1786,  p.  92.  (No  type  locality — Philippine  Islands  ex  Sonnerat,  Voyage 
aux  Indes,  etc.,  1782.) 

Noddy 

Anous  stolidus  (Linnaeus)  in  Rothschild,  Avifauna  of  Laysan,  etc.,  p.  41 ;  Wilson 
and  Evans,  Aves  Hawaiiensis,  p.  141 ;  Bryan,  Key  Birds  Hawaiian  Group, 
p.  9;  Perkins,  Fauna  Hawaiiensis  (Aves),  p.  464;  Henshaw,  Birds  Ha- 
waiian Islands,  p.  124. 


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Range.  Indian  and  western  Pacific  Oceans,  Seychelles  and  Mada- 
gascar, Bonin  and  Hawaiian  Islands  south  to  Australia. 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Laysan 
(breeding),  French  Frigate  Shoal,  Necker,  Bird  Island  (Nihoa), 
Kaula  Lehua  (breeding),  Oahu. 

Anous  minutus  melanogenys  G.  R.  Gray 

Anous  melanogenys  G.  R.  Gray,  Genera  Birds,  3,  1846,  pi.  182.  (No  type 
locality;  figure  agrees  with  Hawaiian  bird  teste  Hartert.) 

Hawaiian  Tern,  Noio 

Anous  hawaiiensis  Rothschild  in  Rothschild,  Avifauna  of  Laysan,  etc.,  vii,  xi, 
43,  285;  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  143;  Perkins,  Fauna 
Hawaiiensis  (Aves),  p.  464;  Henshaw,  Birds  Hawaiian  Islands,  p.  125; 
Microanous  hawaiiensis  Bryan,  Key  Birds  Hawaiian  Group,  p.  9;  et  al. 

Range.     Hawaiian  Islands. 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Laysan 
(breeding),  Lisianski  (breeding),  Necker,  Bird  Rock  (Nihoa),  Lehua, 
Niihau  (sight,  Rothschild),  Kauai,  Oahu  (Mokuloa  and  Kaohi  Kaipu) 
?  Molokai  (W.  K.  Fisher),  Hawaii. 

Genus  Gygis — Fairy  Terns 

Gygis  Wagler,  Isis  von  Oken,  1832,  col.  1223.  Type,  by  monotypy,  Sterna 
Candida  Gmelin. 


Gygis  alba  rothschildi  Hartert 

Gygis  alba  rothschildi  Hartert,  Novitates  Zoologicae,  34,  1927,  p.  18.  (Laysan 
Island.) 

White  Tern,  Love  Bird 

Gygis  alba  (Sparrman)  or  Gygis  Candida  Wagler  in  Rothschild,  Avifauna  of 
Laysan,  etc.,  pp.  vii,  xiii,  xiv,  35,  285;  Wilson  and  Evans,  Aves  Ha- 
waiiensis, p.  145;  Perkins,  Fauna  Hawaiiensis,  p.  145;  Henshaw,  Birds 
Hawaiian  Group,  p.  126;  Gygis  alba  kitllitzi  Hartert  in  Bryan,  Key  Birds 
Hawaiian  Group,  p.  9;  et  al. 

Range.     Islands  northwest  of  Kauai. 

Recorded  in  the  Hawaiian  Group  from  Midway  (breeding),  Laysan 
(breeding),  Lisianski,  French  Frigate  Shoal,  Necker  (breeding), 
Bird  Rock  (Nihoa),  Kaula. 


BRYAN  AND  GREENWAY:  HAWAIIAN   UIRDS  121 

Order  STRIGIFORMES 

Family  STRIGIDAE 

Genus  Asio  Brisson 

Asio  Brisson,  Ornithologie,  1,  1760,  p.  28.  Type,  by  tautonomy,  Asio  Brisson 
=  Strix  otus  Linnaeus. 

Asio  flammeus  sandwichensis  (Bloxam) 

Strix  sandwnchensis  Bloxam,   Voyage   "Blonde",    1826,   p.   250.     (Hawaiian 

Islands.) 
Hawaiian  Owl,  Pueo 
Asio  acciplrinus  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  133;  Perkins, 

Fauna  Hawaiiensis  (Aves),  p.  448. 

Range.  Kauai  (breeding),  Oahu  (breeding),  Molokai  (?  breeding), 
Maui,  Hawaii  (breeding). 

Order  CORACIIFORMES 

Family  ALCEDINIDAE 

Genus  Ceryle — Kingfishers 

Ceryle  Boie,  Isis  von  Oken,  1828,  col.  316.  Type,  by  subsequent  designation , 
Alcedo  rudis  Linnaeus,  Gray,  List  Genera  Birds,  1840,  p.  11. 

Ceryle  alcyon  caurina  Grinnell 

Ceryle  alcyon  caurina  Grinnell,  University  California  Publications,  Zool.,  6> 
1910,  p.  388.  (Graveyard  Point,  Montague  Island,  Prince  William 
Sound,  Alaska.) 

Western  Belted  Kingfisher 

Ceryle  alcyon  (Linnaeus)  in  Henshaw,  Birds  Hawaiian  Group,  p.  77. 

Range.  Alaska  south  to  southern  California;  south  to  northern 
and  central  Mexico  in  winter. 

Recorded  in  the  Hawaiian  Group  (as  an  accidental  visitor)  from 
Hawaii. 

Order  PASSERIFORMES— Perching  Birds 

Family  CORVIDAE 

Genus  Corvus — Crows 

Corvus  Linnaeus,  Systema  Naturae,  ed.  10,  1,  1758,  p.  105.  Type,  by  Linnaean 
tantonymy,  Corvus  corax  Linnaeus. 


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Corvus  hawaiiensis  Peale 

Corvus  hawaiiensis  Peale,  U.  S.  Exploring  Expedition,  8,  1848,  p.  106.  (Kara- 
kakua  [Kealakekua]  Bay,  Hawaii.) 

Hawaiian  Crow,  Alala 

Corvus  tropicus  "Linnaeua",  Bloxam  in  Voyage  "Bonde",  1826,  p.  250  (Sand- 
wich Islands)  is  a  nomen  nudum.  Corvus  tropicus  Gmelin  to  which  Bloxam 
refers  is  not  identifiable. 

Corvus  tropicus  Gm.  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  1 ;  Henshaw, 
Birds  Hawaiian  Possessions,  etc.,  p.  36. 

Range.     Edges  of  mountain  forest  from  1000  to  6000  feet  in  Kona 
and  Kau  districts  of  the  Island  of  Hawaii. 


Family  MUSCICAPIDAE— Flycatchers,  Thrushes,  Warblers 
Subfamily  TURDINAE 

Genus  Phaeornis — Hawaiian  Thrushes 

Phaeornis  Sclater,  Ibis,  1,  1859,  p.  327.    Type,  by  original  designation  and 
monotypy,  Phaeornis  obscura  (Gmelin). 

Phaeornis  obscura  obscura  (Gmelin) 

Muscicapa  obscura  Gmelin,  Systema  Naturae,   1,  pt.   2,    1789,  p.  945.   "(in 

insulis  Sandwich.") 
Ornao,  Oman,  Hawaii  Thrush,  Kamao,  A-Maui. 

Range.     Endemic  to  Island  of  Hawaii. 

Phaeornis  obscura  lanaiensis  Wilson 

Phaeornis  lanaiensis  Wilson,  Annals  and  Magazine  Nat.  Hist.  (6),  7,  1891 » 

p.  460.    (Lanai.) 
Lanai  Thrush,  OXomao,  Olomau 

Range.     Mountain  forests  of  Lanai.   Now  very  rare. 

Phaeornis  obscura  rutha  W.  A.  Bryan 

Phaeornis  rutha  W.  A.  Bryan,  Occasional  Papers  B.  P.  Bishop  Mus.,  Honolulu, 

4,  no.  2,  1908,  p.  81.    (Molokai.) 
Molokai  Thrush,  Olomao 

Range.     Mountain  forests  of  Molokai. 

This  poorly  marked  form  averages  somewhat  larger  (wing  averages 
92  mm.  and  95  mm.)  and  slightly  darker  than  lanaiensis. 


BRYAN  AND  GRKKNWAV:  HAWAIIAN  BIRDS  123 

Phaeornis  ?  obscurus  oahensis  Wilson  and  Evans 

Phaeornis   oahensis    Wilson    and    Evans,    Aves    Hawaiiensis    (Introduction), 

1899,  p.  XIII  (Oahu). 
Oahu  Thru  ah 

Range.     Formerly  mountain  forests  of  Oahu.   Now  extinct. 

Specimens  are  supposed  to  have  been  collected  by  Andrew  Bloxam, 
naturalist  on  H.  M.  S.  'Blonde',  but  they  have  disappeared.  Wilson 
and  Evans  described  the  bird  from  Bloxam's  manuscript  notes. 


Phaeornis  obscura  myadestina  Stejneger 

Phaeornis  myadestina  Stejneger,  Proceed.  U.  S.  Nat.  Mus.,  10,  1887  (1888), 
p.  90.    (Kauai.) 

Phaeornis  myiadestina  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  117;  Roths- 
child, Avifauna  Laysan,  p.  63. 

Kauai  Thrush,  Kamau,  Kamao 

Range.     Forests  of  Kauai. 

Phaeornis  palmeri  Rothschild 

Phaeornis  palmeri  Rothschild,  Avifauna  Laysan  Island,  etc.,  pt.  2,  1893,  p.  67. 

(Halemanu,  Kauai.) 
Puaiohi 

Range.  Region  of  Halemanu,  Kauai.  Never  recorded  from  other 
parts  of  the  island.   Now  very  rare  and  possibly  extinct. 

This  bird,  because  of  its  voice  and  habits,  which  differ  from  those 
of  other  members  of  the  genus  (fide  Perkins  et  al.),  and  because  it 
apparently  breeds  in  the  same  region  as  myadestina,  we  must  treat 
as  a  distinct  species. 


Subfamily  SYLVIINAE 

Genus  Acrocephalus- — Miller  Birds 

Acrocephalus  Naumann,  Land  u.  Wasservogel  nordl.  Deutschland,  Nachtrage, 
4,  1811,  p.  199.  Type,  by  subsequent  designation,  Sylvia  turdoides  Meyer 
=  Acrocephalus  arundinaceus  (Linnaeus). 

We  can  see  no  reason  for  retaining  the  genus  Conopoderas  Billberg. 
Some  of  the  island  forms  are  closer  to  the  type  of  Acrocephalus  than 
to  that  of  Conopoderas. 


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ACROCEPHALUS  FAMILIARIS  (Rothschild) 

Tatare  familiaris  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10,  1892, 

p.  109.    (Laysan  Island.) 
Laysan  Miller  Bird 

Range.     Formerly  Laysan  Island,  now  extinct. 


Acrocephalus  kingi  ( Wetmore) 

Conopoderas  kingi  Wetmore,  Condor,  26,  1924,  p.  177.    (Nihoa.) 
Nihoa  Miller  Bird 

Range.     Nihoa  (Bird  Rock). 

In  employing  binomials  we  do  not  mean  to  imply  that  these  forms 
are  distinct  species.  The  present  application  of  specific  names  is 
unsatisfactory.   This  genus  needs  revision. 


Subfamily  MUSCICAPINAE 

Genus  Chasiempis  Cabanis 

Chasiempis  Cabanis,  Archiv  fur  Naturgeschiehte,  13,  1847,  Bd.  1,  p.  207. 
Type,  by  original  designation,  "Muscicapa  sandvicensis  Latham"  = 
Chasiempis  sandwichensis  (Gmelin). 

Though  Cabanis  cited  Muscicapa  sandvicensis  Latham,  Latham 
himself  never  used  this  name.  Gmelin 's  description  of  Muscicapa 
sandwichensis  was  taken  from  Latham's  "Sandwich  Fly-catcher." 


Chasiempis  sandwichensis  sclateri  Ridgway 

Chasiempis  sclateri  Ridgway,  Proceed.  U.  S.  Nat.  Mus.,  4,  1881,  (1882),  p.  337. 

(Waimea,  Kauai.) 
Kauai  Elepaio,  Apekepeke,  Amakahi,  Kahuna-Ka-lai-woa 
Chasiempis  dolei  Stejneger,  Proceed.  U.  S.  Nat.  Mus.,  10,  1887  (1888),  p.  90. 

(Kauai.) 

Range.     Forests  of  Kauai. 

Both  Hawaiians  and  scientists  were  long  under  the  impression  that 
the  white-rumped  form  (i\makahi  or  Elepaio)  and  the  brown-rumped 
form  (Apekepeke)  were  distinct  species.  Perkins  showed  that  the 
latter  is  an  immature  plumage  in  which  the  birds  sometimes  breed. 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  125 


Chasiempis  sandwichensis  gayi  Wilson 

Chasiempis  gayi  Wilson,  Proceed.  Zool.  Soc.  London,  1891,  p.  165.    (Oahu.) 
Oahu  Elepaio 


Range.     Forests  of  Oahu. 


Chasiempis  sandwichensis  sandwichensis  (Gmelin) 

Muscicapa  sandwichensis  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  945. 

("in  insulis  Sandwich"  =  Hawaii.) 
Hawaii  Elepaio,  Ono-Ka-Ia 
Chasiempis  rirfgwayi  Stejneger,  Proceed.  U.  S.  Nat.  Mus.,  10,  1887  (1888), 

p.  87,  in  key,  p.  89.    (Hawaii.) 
Chasiempis  ibidis  Stejneger,  Proceed.  U.  S.  Nat.  Mus.,  10,  1887  (1888),  p.  87, 

in  key,  p.  89. 

Range.     Forests  of  Hawaii. 

Stejneger's  Chasiempis  ibidis  refers  to  plate  1,  figure  2  in  the  Ibis 
of  1S85.  This  specimen  had  no  locality  other  than  "Chili,"  as  Sclater 
remarks. 


Subfamily  DREPANIDIXAE 

Genus  Viridonia 

Yiridonia  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10,  1892,  p.  112. 
Type,  by  monotypy,  Viridonia  sagittirostris  Rothschild. 


Viridonia  sagittirostris  Rothschild 

Viridonia  sagittirostris  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10, 

1892,  p.  112.    (Lower  Hilo  slope  of  Mauna  Kea,  Hawaii.) 
Green  Solitaire 

Range.     Mountain  rain  forests  of  the  windward  side  of  Hawaii  near 
the  Wailuku  River  at  about  1200  to  4000  feet. 


Genus  Palmeria 

Palmeria  Rothschild,  Ibis,  1893,  p.  113.    Type,  by  original  designation  and 
monotypy,  Palmeria  mirabilis  Rothschild  =  P.  dolei  I  Wilson). 


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Palmeria  dolei  (Wilson) 

Himatione  dolei  Wilson,  Proceed.  Zool.  Soc.  London,  1891,  p.  166.    (Maui.) 

Akohekohe,  Crested  honey  eater 

Palmeria  mirabilis  Rothschild,  Ibis,  1893,  p.  113. 

Range.     Mountain  forests  of  Molokai  and  Maui.  Now  very  rare. 

Genus  Himatione 

Himatione  Cabanis,  Museum  Heineanum,  Th.  1,  1850,  p.  99.  Type,  by  subse- 
quent designation,  Himatione  sanguinea  (Gmelin). 

Himatione  sanguinea  fraithii  Rothschild 

Himatione  Fraithii  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10,  1892, 

p.  109.    (Laysan  Island.) 
Laysan  Honey  Creeper 

Range.     Formerly  Laysan  Island,  now  extinct. 

Himatione  sanguinea  sanguinea  (Gmelin) 

Certhia  sanguinea  Gmelin,  Systema  Naturae,  1,  pt.  1,  1788,  p.  479.    (Sandwich 

Islands.) 
Apapane,  Akakani 

Range.     Mountain    forests    of    ?Niihau,    Kauai,    Oahu,    Molokai, 
Maui,  Hawaii. 

Genus  Vestiaria 

Vestiaria  Fleming,  Philosophy  of  Zoology,  2,  1822,  p.  246.   Type,  by  original 
designation,  Certhia  vestiaria  =  Vestiaria  coccinea  (Forster). 


Vestiaria  coccinea  coccinea  (Forster) 

Certhia  coccinea  Forster,   Goettingisches   Magazin   Wissenschaften,   etc.,    1, 

1780,  p.  347. 
Iiwi,  Iiwi  popolo,  Iiwi  polena  (for  various  phases  of  plumage). 
Vestiaria  coccinea  suavis  Bangs,  Proceed.  Biol.  Soc.  Wash.,  24,  1911,  p.  29. 

(Molokai.) 

Range.     Mountain  forests  of  Kauai,  Oahu,  Molokai,  Maui,  Lanai, 
Hawaii. 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  127 

Tor  full  synonymy  see  Rothschild,  Avifauna  Laysan  Island,  etc. 

Vestiaria  coccinea  suavis  Bangs  was  described  as  paler,  with  a  larger 
bill.  In  our  opinion,  the  former  is  an  individual  variation  and  the  latter 
is  non-existent.  In  our  short  series  (28)  there  is  an  average  difference 
of  2  mm.  in  wing  length,  but  this  is  not  sufficient  for  formal  recognition. 

Genus  Drepaxis — Hawaiian  Honey  Creepers 

Drepanis  Temminck,  Manual  d'Ornithologie,  ed.  2,  1,   1820,  p.  LXXXVI . 
Type,  by  subsequent  designation  (Gray,  1840),  Certhia  pacifica  Gmelin. 

Drepaxis  pacifica  (Gmelin) 

Certhia  pacifica  Gm.,  Systema  Naturae,  1,  pt.  1,  1788,  p.  470.    ("in  insulis 

amicis"  in  error  ex  Latham.   Hawaiian  Islands.) 
Mamo,  Hoha  or  Hoho  (so  rendered  by  older  writers). 

Range.  Formerly  mountain  forests  of  Hawaii  (old  records  for 
Kauai  are  almost  certainly  in  error).  Now  extinct;  last  specimen  seen 
at  Kaumana  1000-1500  feet  above  Hilo,  1898. 

Drepaxis  fuxerea  Newton 

Drepanis  funerea  Newton,  Proceed.  Zool.  Soc.  London,  p.  690.  (Molokai.) 

Ma?no,  Oo-nuku-umu,  Hoa,  Black  Mamo,  Perkins'  Mamo 

Drepanorhamphus  funerea  Newton  in  Rothschild,  Avifauna  Laysan  Island, 

etc.,  p.  165;  Bryan,  Birds  Hawaiian  Group,  p.  42;  Perkins,  Fauna  Ha- 

waiiensis  (Avesj,  p.  402. 

Range.     Mountain  forests  of  Molokai. 

The  genus  Drepanorhamphus  was  established  by  Rothschild  (Avi- 
fauna Laysan,  etc.,  pt.  3,  1900,  p.  163.  Type,  by  monotypy,  Drepanis 
funerea  Newton).  The  characters  that  separate  it  from  Drepanis  are 
the  lack  of  the  long,  decomposed,  yellow  under  tail  coverts  and  the 
shape  of  the  nostril  which  is  long  and  narrow,  not  rounded.  It  seems 
to  us  that  they  are  alike  in  so  many  ways  that  they  form  a  single  genus 
separating  them  from  other  genera,  but  that  they  were  probably  dis- 
tinct species. 

Genus  Hemigxathus — Akialoa 

Hemignathus  Lichtenstein,  Abhandlungen  Konig.  Akademie  Wissensch 
Berlin,  1839,  p.  449.  Type,  by  subsequent  designation,  Hemignathus 
lucidus  Licht.    (Gray,  List  Genera  Birds,  1841,  p.  16). 


128  bulletin:  museum  of  comparative  zoology 

In  our  opinion,  the  group  with  long  lower  mandibles  and  those  with 
short  lower  mandibles  may  well  be  considered  as  congeneric.  If  it  is 
desired  to  separate  them  generically  then  the  former  will  require  a 
new  generic  name  and  the  latter  (heretofore  known  as  Heterorhynchus 
will  have  to  be  called  Hemignathus,  since  the  two  groups,  as  named 
heretofore,  have  the  same  type  as  designated  by  Gray  and  Lafresnaye. 
Rothschild's  arguments  (Avif.  Laysan  Id.,  etc.,  p.  79)  have  no  force 
under  the  rules  of  zoological  nomenclature  (Art.  30,  II,  g). 

Hemignathus  obscurus  procerus  Cabanis 

Hemignathus  procerus  Cabanis,  Journal  f.  Ornith.,  1889,  p.  331.    (Kauai.) 
Kauai  Akialoa,  "Iiwi" 

Range.  Mountain  forests  of  Kauai  from  the  lowest  forest  zone 
600-900  feet  to  4000  feet. 

Hemignathus  obscurus  ellisianus  (Gray) 

Drepanis  {Hemignathus)  ellisiana  Gray,  Cat.  Birds  Tropical  Islands  Pacific, 

1860,  p.  9.    (Oahu.) 
Oa.hu  Akialoa,  Iiwi  (Jibi  or  Kipi) 
Hemignathus  lichtensteini  Wilson  in  Aves  Hawaiiensis,    p.  65,  and  Perkins, 

Fauna  Hawaiiensis  (Aves),  p.  425. 

Range.  Formerly  mountain  forests  of  Oahu;  now  extremely  rare; 
perhaps  confined  to  Waianae  Mountains  (Northwood). 

Hemignathus  obscurus  lanaiensis  Rothschild 

Hemignathus  lanaiensis  Rothschild,  Bull.  Brit.  Ornith.  Club,  1,  1893,  p.  xxiv. 

(Lanai.) 
Lanai  Akialoa 

Range.  Formerly  mountain  forests  of  Lanai,  now  very  rare,  pos- 
sibly extinct. 

Hemignathus  obscurus  obscurus  (Gmelin) 

Certhia  obscura  Gm.,  Systema  Naturae,  1,  pt,  1,  1788,  p.  470.    (Sandwich 

Islands  =  Hawaii.) 
Haivaii  Akialoa 

Range.  Formerly  mountain  forests  of  Hawaii,  now  probably  con- 
fined to  localized  areas  in  the  rain  forests  of  the  windward  side  (see 
Baldwin,  1941,  p.  19). 


BRYAN  AND  (iKKF.NW  AY:  HAWAIIAN  BIRDS  129 


Hemignathus  lucidus  iiaxapepe  Wilson 

Hemignathus  hanapepe  Wilson,   Annals  Magazine  Nat.  Hist.   (6),  4,   1889, 

p.  401.    (Kauai.) 
Nukupu'u 
Heterorhgnrhus  hanapt  /«  in  Rothschild,  Avifauna  Laysan,  etc.,  p.  101 ;  Perkins, 

Fauna  Hawaiiensis  (Aves),  p.  430;  Henshaw,  Birds  Hawaiian  Islands, 

p.  43. 

Range.     Mountain  forests  of  Kauai  in  Waimea  district  near  the 
Hanapepe  River,  2000  to  3000  feet,  very  rare  and  local. 


Hemignathus  lucidus  lucidus  Lichtenstein 

Hemignathus  lucidus  Lichtenstein,  Abhandlungen  der  Konigl.  Academie 
Wissensch.  Berlin,  1S39,  p.  451,  pi.  5,  figs.  2,  3.    (Oahu.) 

Oahu  Akiapolaau 

HeterorhyncJms  lucidus  in  Rothschild  Avifauna  Laysan,  etc.,  p.  105;  Perkins, 
Fauna  Hawaiiensis  (Aves),  p.  430;  Henshaw,  Birds  Hawaiian  Islands, 
p.  41. 

Akiapolaau 

Range.  Formerly  mountain  forests  of  Oahu ;  now  almost  certainly 
extinct. 

Ten  specimens  were  thought  to  exist  in  museums  but  there  are  not 
that  many,  for  some  have  been  misidentified. 


Hemignathus  lucidus  affinis  Rothschild 

Hemignathus  affinis  Rothschild,  Ibis,  1893,  p.  112.    (Maui.) 
Maui  Akiapolaau 

Range.     Recorded  from  forested  slopes  of  Haleakala  Mountain, 
Maui,  4000  to  4500  feet.  Now  local  and  probably  very  rare. 


Hemignathus  lucidus  wilsoni  (Rothschild) 

Heterorhynchus  wilsoni  Roths.,  Avifauna  Laysan,  etc.,  1893,  pt.  II,  p.  97. 

(Hawaii). 
Akiapolaau 
Hemignathus  olivaceus  Lafr.  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  75. 

Range.     Mountain  forests  of  Hawaii  from  about  1500  to  6700  feet. 
Still  exists  but  not  common. 


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Genus  Chlorodrepanis — Amakihi 

Chlorodrepanis  Perkins  in  Wilson  and  Evans,  Aves  Hawaiiensis,  pt.  7,  p.  xxi 
(introduction),  June  1899.  Type,  by  subsequent  designation,  Chlorodre- 
panis stejriegeri  Wilson  (Richmond,  Proceed.  U.  S.  Nat.  Mus.,  24,  1902, 
p.  673). 


Chlorodrepanis  parva  (Stejneger) 

Himatione  parva  Stejneger,  Proceedings  U.  S.  Nat.  Mus.,  10,  1887,  p.  94. 

(Kauai.) 
Alawi,  Anauanii,  Anianau 
Oreomyza  parva  in  Rothschild,  Avifauna  Laysan,  etc.,  p.  119. 

Range.     Forests  of  Kauai. 

This  is  a  very  curious  little  bird  for  which  Wilson  and  Evans  pro- 
posed the  name  Rothschildia  (pre-occupied).  Mathews  in  1925  re- 
named it  Magumma.  Our  treatment  is  based  on  that  of  Perkins 
(Ibis,  1895  and  Fauna  Hawaiiensis  (Aves),  p.  411). 


Chlorodrepanis  virens  stejnegeri  (Wilson) 

Himatione  stejnegeri  Wilson,  Proceedings  Zool.  Soc.  London,   1889   (1890), 

p.  446.    (Kauai.) 
Kauai  Amakihi 

Range.     Forests  of  Kauai. 

Chlorodrepanis  virens  wilsoni  (Rothschild) 

Himatione  wilsoni  Rothsch.,  Bull.  Brit.  Ornith.  Club,  1,  1893,  p.  xliii.    (Maui.) 

Molokai  Amakihi,  Lanai  Amakihi,  Maui  Amakihi 

Himatione  kalaana  Wilson  and  Evans,  Aves  Hawaiiensis,  1896,  p.  28.  (Molo- 
kai.) 

Himatione  chloridoides  Wilson  and  Evans,  Aves  Hawaiiensis,  1896,  p.  28. 
(Lanai.) 

Range.     Forests  of  Molokai,  Lanai,  Maui. 

This  form  is  very  close  to  chloris  but  differs  in  having  a  slightly 
larger  bill  and  (in  mature  specimens)  it  has  a  slightly  paler  back. 
Differences  between  specimens  from  Molokai,  Lanai  and  Maui  alleged 
to  exist  (an  ill-defined  or  well-defined  yellow  stripe  from  bill  to  eye  as 
well  as  the  shade  of  green  on  the  back)  fall  within  the  range  of  indi- 
vidual variation. 


BRYAN  AND  GREENWAT:  HAWAIIAN  BIRDS  131 


Chlorodrepanis  virens  virens  (Gmelin) 

Certhia  virens  Gm.,  S}'stema  Naturae,  1,  pt.  1,  17S8,  p.  479.    (in  insulis  Sand- 
wich =  Hawaii.) 
Hawaii  Amakihi 

Range.     Forests  of  Hawaii. 

Genus  Loxops — Akakane 

Loxops  Cabanis,  Archiv  fur  Naturgeschichte,  Berlin,  13,  1847,  Bd.  1,  p.  330. 
Type,  by  original'  designation,  Fringilla  coccinea  Gm.  For  synonymy  see 
Rothschild,  Avifauna  Laysan,  etc.,  p.  167. 


Loxops  caeruleirostris  (Wilson) 

Chrysomitridops  caeruleirostris  Wilson,  Proceedings  Zool.  Soc.  London,  1889 

(1890),  p.  445.    (Kauai.) 
Ou-holoicai,  Akekee  (fide  Perkins) 
Chrysomitridops  caeruleirostris  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  59. 

Range.     Forests  of  Kauai. 

Loxops  coccinea  rufa  (Bloxam) 

Fringilla  rufa  Bloxam  in  Voyage  "Blonde",  1826,  p.  250.    (Oahu.) 
Akepa,  Akepeuie 

"Loxops  wolstenkolmei"  Roths. — the  plate  so  lettered  in  Rothschild  Avifauna 
Laysan,  etc.,  p.  177. 

Range.     Formerly  forests  of  Oahu,  now  very  rare  and  perhaps 
extinct.   Last  record  May  20,  1S93. 

Loxops  coccinea  ochracea  Rothschild 

Loxops  ochracea  Rothschild,  Ibis,  1893,  p.  112.    (Maui.) 
Ochraceus  or  Maui  Akt  peuie 

Range.     Forests  of  Maui. 

Loxops  coccinea  coccinea  (Gmelin) 

Fringilla  coccinea  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  921.    (in  insulis 

Sandwich  =  Hawaii.) 
Akepa,  Akepeuie 

Range.     Forests  of  Hawaii.    "Now  rare"  (Baldwin). 


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Genus  Paroreomyza 

Paroreomyza  Perkins,   Ibis,    1901,   p.    583.     Type,   by   original   designation, 
Himatione  maculata  Cabanis. 

In  our  opinion,  the  forms  formerly  listed  under  this  name  and  those 
listed  under  the  name  Oreomystis  Stejneger  1903  are  congeneric.  It  is 
true  that  females  of  the  former  lack  the  color  of  the  males,  but  the 
birds  are  otherwise  alike  from  a  generic  point  of  view. 


Paroreomyza  bairdi  bairdi  (Stejneger) 

Oreomyza  bairdi  Stejneger,  Proceedings  U.  S.  Nat.  Mus.,  10,  1887  (1888),  p.  99. 

(Kauai.) 
Akikihi 
Oreomyza  bairdi  in  all  works  on  Hawaiian  birds. 

Range.     Forests  of  Kauai  above  1000  feet. 

Paroreomyza  bairdi  mana  (Wilson) 

Himatione  mana  Wilson,  Annals  and  Magazine  Nat.  Hist.  (6),  7,  1891,  p.  460. 

(Hawaii.) 
Hawaii  Creeper,  Olive-green  Creeper 
Oreomyza  mana  in  all  works  on  Hawaiian  birds. 

Range.  Forests  of  Hawaii  except  (fide  Perkins)  in  lower  elevations 
on  the  Kona  or  leeward  coast.   Now  uncommon  (fide  Baldwin  1941). 

In  our  opinion  "Oreomyza  perkinsi"  Rothschild  is  in  all  probability 
a  hybrid,  Chlorodrepanis  virens  x  Paroreomyza  mana.  It  has  been  listed 
with  other  hypothetical  forms  and  records. 

Paroreomyza  maculata  maculata  (Cabanis) 

Himatione  maculata  Cabanis,  Museum  Heineanum,  Th.   1,  1850,  p.   100,  in 

footnote.    (Oahu.) 
Oahu  Creeper 
Himatione  maculata  Cabanis  in  Wilson  and  Evans,  Aves  Hawaiensis,  p.  43; 

Oreomyza  maculata  in  Rothschild,  Avifauna  Laysan  Island,  etc.,  p.  113; 

W.  A.  Bryan,  Key  Birds  Hawaiian  Group,  p.  48;  Perkins,  Fauna  Ha- 

waiiensis,  (Aves)  p.  417;  Henshaw,  Birds  Hawaiian  Islands,  p.  50. 

Range.  Mountain  forests  of  Oahu  above  1500  feet.  "Scarcest  of 
the  five  native  perching  birds  which  are  likely  to  be  seen."  (North- 
wood  1940). 


BRYAN  AND  OKEENWAY :  HAWAIIAN   BIRDS  133 


Paroreomyza  maculata  flammea  (Wilson) 

Loxops  flarnmea  Wilson,  Proceedings  Zool.  Soc.  London,  1889  (1890),  p.  445. 
(Kalae,  Molokai.) 

Kakawahie 

Loxops  flammea  Wilson  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  39;  Oreomyza 
flammea  Wilson  in  Rothschild,  Avifauna  Laysan  Island,  etc.,  p.  121; 
W.  A.  Bryan,  Key  Birds  Hawaiian  Group,  p.  48;  Perkins,  Fauna  Hawaii- 
ensis (Aves),  p.  417;  Henshaw,  Birds  Hawaiian  Islands,  p.  49. 

Range.     Mountain  forests  of  Molokai  above  1500  feet. 


Paroreomyza  maculata  Montana  (Wilson) 

Himatione  montana  Wilson,  Proceedings  Zool.  Soc.  London,  1889  (1890),  p. 
446.    (Lanai.) 

Alauhiio 

Himatione  montana  Wilson  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  45; 
Oreomyza  montana  in  Rothschild,  Avifauna  Laysan  Island,  etc.,  p.  117; 
W.  A.  Bryan,  Key  Birds  Hawaiian  Group,  p.  47;  Perkins,  Fauna  Hawaii- 
ensis (Aves),  p.  417;  Henshaw,  Birds  Hawaiian  Islands,  p.  51. 

Range.     Mountain  forests  of  Lanai. 


Paroreomyza  maculata  newtoni  (Rothschild) 

Himatione  newtoni  Rothschild,  Bull.  British  Ornith.  Club,  1,  1893,  p.  xlii. 

(Maui.) 
Maui  Creeper 
Himatione  newtoni  Rothschild  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  11; 

Oreomyza  newtoni  Rothschild  in  Rothschild,  Avifauna  Laysan  Island,  etc. 

p.  115;  W.  A.  Bryan,  Key  Birds  Hawaiian  Group,  p.  48;  Perkins,  Fauna 

Hawaiiensis  (Aves),  p.  417;  Henshaw,  Birds  Hawaiian  Islands,  p.  49. 

Range.     Mountain  forests  of  Maui. 


Genus  Ciridops 

Ciridops  A.  Newton,  Nature,  45,  1892,  p.  469.   Type,  by  monotypy,  Fringilla 
anna  Dole. 


Ciridops  anna  (Dole) 

Fringilla  anna  Dole,  Hawaiian  Almanac,  1879,  p.  49.    (Hawaii.) 
Waaihaiuane 


134  bulletin:  museum  of  comparative  zoology 

Range.  Formerly  mountain  forests  of  Hawaii.  Recorded  from 
Kona  and  Hilo  districts  in  or  near  Loulu  or  Hawane  palms  (Prit- 
chardia).  Never  common  within  the  memory  of  man,  it  is  now  prob- 
ably extinct.  Last  specimen  taken  February  1892  on  the  Kohala 
Mountains. 


Genus  Pseudonestor 

Pseudonestor  Rothschild,   Bulletin   British  Ornith.   Club,   1,    1893,   p.   xxxv. 
Type,  by  monotypy,  Pseudonestor  xanthophrys  Rothschild. 


Pseudonestor  xanthophrys  Rothschild 

Pseudonestor  xanthophrys  Rothschild,  Bulletin  British  Ornith.  Club,  1,  1893, 

p.  36.    (Maui.) 
ParroUbilled  Koa  Finch 

Range.     Maui;  confined  to  Haleakala  above  5000  feet.   Rare. 


Genus  Psittirostra 

Psittirostra  Temminck,  Manuel  d'Ornithologie,  etc.',  1,  1820,  p.  70.   Type,  by 
monotypy,  Loxia  psittacea  Gmelin. 

This  name  has  often  been  emended  to  Psittacirostra;  the  original 
spelling  is,  however,  as  above. 


Psittirostra  psittacea  psittacea  (Gmelin) 

Loxia  psittacea  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  844.  (in  insulis 
Sandwich  =  Hawaii.) 

Ou 

Psittacirostra  psittacea  oppidana  Bangs,  Proceedings  Biol.  Soc.  Washington, 
24,  1911,  p.  30.    (Molokai.) 

Examination  of  large  series  has  convinced  us  that  characters  alleged  to  differen- 
tiate this  form  from  psittacea  (paleness,  size)  fall  within  the  range  of 
individual  variation. 

Dysmorodrepanis  munroi  Perkins  (see  list  of  hypothetical  forms). 

Range.     Forests  of  Molokai,  Lanai,  Maui,  Hawaii  below  5000  feet. 
Now  very  rare,  possibly  extinct  on  Lanai. 


BRYAN  AND  GREEXWAY  :  HAWAIIAN'  BIRDS  135 


?PsiTTIR0STRA  PSITTACEA  DEPPEI  Rothschild 

PsiUirostra  psittacea  deppei  Rothschild,  Bull.  Brit.  Ornith.  Club,  15,  1905-, 
p.  45.    (Oahu.) 

Oahu  Ou 

Psittacirostra  psittacea  (Gmelin)  (part)  in  Wilson  and  Evans,  Aves  Hawaiiensis, 
p.  80;  PsiUirostra  olivacea  Rothschild  in  Rothschild,  Avifauna  Laysan 
Island,  etc.,  p.  193;  W.  A.  Bryan,  Key  Birds  Hawaiian  Group,  p.  54 
(note);  Henshaw,  Birds  Hawaiian  Islands,  p.  66;  Perkins,  Fauna  Ha- 
waiiensis (Aves),  p.  435. 

Range.     Formerly  forests  of  Oahu,  now  extinct. 

There  is  great  individual  variation  in  adults  of  the  populations  of  this 
bird  on  every  island.  Particularly,  there  is  a  tendency  toward  albinism 
noticeable  in  small  populations.  Rothschild  describes  this  form  as 
differing  from  others  in  having  the  middle  of  the  breast,  belly, feathers 
of  the  tibia  and  under  tail  coverts  buffy  whitish,  in  having  a  shorter 
wing.  In  view  of  the  fact  of  this  known  tendency  toward  albinism  and 
that  Rothschild's  own  measurement  for  the  wing  of  deppei  (3.75 
inches  =  97  mm.)  falls  exactly  within  the  size  range  of  psittaeea 
(95-100  mm.),  we  think  the  validity  of  this  subspecies  is  questionable. 


Genus  Loxioides — Palila 

Loxioides  Oustalet,  Bulletin  Soc.  Philom.  Paris,  ser.  7,  1,  1877,  p.  99.  Type,  by 
monotypy,  Loxioides  bailleui  Oustalet. 


Loxioides  bailleui  Oustalet 

Loxioides  bailleui  Oustalet,  Bulletin  Soc.  Philom.  Paris,  ser.  7,  1,  1877,  p.  100. 

(Hawaii.) 
Palila 
Loxioides  bailleui  Oustalet  in  all  works  relating  to  Hawaiian  birds. 

Range.     Upper  forest  zones  of  Hawaii  4000  to  7000  feet  in  Kona  and 
Hamakua  districts.   Usually  in  or  near  mamane  trees  (Sophora). 


Genus  Rhodacanthis — Koa  Finch 

Rhodacanthis  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10,  1892,  p. 
110.   Type,  by  subsequent  designation,  Rhodacanthis  palmeri  Rothschild. 


136  bulletin:  museum  of  comparative  zoology 

Rhodacanthis  palmeri  Rothschild 

Rhodacanthis  Palmeri  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10, 

1892,  p.  111.    (Kona,  Hawaii.) 
Orange  Koa  Finch 

Range.  Above  4000  feet  in  mountain  forests  of  Kona  and  Kau 
districts,  Hawaii.  Usually  in  Koa  trees  (Acacia  Koa).  Now  very  rare, 
possibly  extinct.   Last  specimens  collected  (1892  or  1893)  by  Perkins. 

?Rhodacanthis  flaviceps  Rothschild 

Rhodacanthis  flaviceps  Rothschild,  Annals  and  Magazine  Nat.  Hist.  (6),  10' 

1892,  p.  111.    (Kona,  Hawaii.) 
Yellow-headed  Koa  Finch 

Range.  Presumably  the  same  as  Rhodacanthis  palmeri  with  which 
it  was  associated  when  collected  by  Palmer  for  Rothschild. 

These  specimens  differ  in  having  yellow,  not  red,  heads  and  in  being 
smaller  (wing  94-95  mm.).  No  specimens  have  since  been  found. 
Perkins  (Fauna  Hawaiiensis  (Aves),  pp.  436-437)  suggests  that  these 
are  not  two  distinct  species  but  that  only  one  dimorphic  form  is  in- 
volved. Since  the  genus  is  now  very  rare  it  is  probable  that  this 
problem  can  never  be  solved. 

Genus  Telespyza 

Telespyza  Wilson,  Ibis,  1890,  p.  341.    Type,  by  monotypy,  Telespyza  cantans 
Wilson. 

Telespyza  cantans  cantans  Wilson 

Telespyza  cantans  Wilson,  Ibis,  1890,  p.  341.    (Midway  Island  in  error  = 

Laysan  Island.) 
Lay san  Finch 
Telespyza  flavissima  Rothschild,  Annals  and  Magazine  Nat.  Hist.   (6),   10, 

1892,  p.  110.    (Laysan.) 

Range.  Formerly  on  Laysan  Island,  where  now  extinct.  Midway 
Island,  where  introduced. 

Telespyza  cantans  ultima  W.  A.  Bryan 

Telespiza  ultima  W.  A.  Bryan,  Auk,  34,  1917,  p.  71.    (Nihoa.) 
Nihoa  Finch 

Range.     Nihoa. 


BRYAN  AND  GREEXWAY:  HAWAIIAN  BIRDS  137 

Genus  Chloridops 

Chloridops  Wilson,  Proceedings  Zool.  Soc.  London,  1888,  p.  218.    Type,  by 
monotypy,  Chloridops  kona  Wilson. 

Chloridops  kona  Wilson 

Chloridops  kona  Wilson,  Proceedings  Zool.  Soc.  London,  1888,  p.  218.    (Kona, 

Hawaii.) 
Kona  Finch,  ?  Palila 

Range.     Kona  district  of  Hawaii,  in  mountain  forests  from  3500  to 
5500  feet.   Never  abundant,  now  very  rare,  perhaps  extinct. 


Family  MELIPHAGIDAE 
Genus  Moho 

Moho  Lesson,  Traite  d'Ornithologie,  livr.  4,  1830,  p.  302.  Type,  by  monotypy, 
Merops  fasciculatus  Latham  =  Moho  nobilis  Merrem. 

Mohohina  Mathews,  Bulletin  British  Ornith.  Club,  45,  1925,  p.  93.  Type,  by 
original  designation,  Acridocercus  bishopi  Rothschild.  "Differs  from 
Moho  Lesson  in  having  plumes  on  the  side  of  the  face"  (Mathews). 

Pseudomoho  Mathews,  Bulletin  British  Ornith.  Club,  45,  1925,  p.  93.  Type,  by 
original  designation,  Mohoa  braccatus  Cassin.  "Differs  from  Moho  Lesson 
in  having  quite  a  different  tail  formation  and  in  not  having  any  orna- 
mental plumes"  (Mathews). 

Mohornis  Mathews,  Systema  Avium  Australasianarum,  pt.  2,  1930,  p.  800. 
Type,  by  original  designation,  Moho  apicalis  Gould. 

Moho  nobilis  braccatus  (Cassin) 

Mohoa  braccata  Cassin,  Proceedings  Acad.  Nat.  Sci.  Philadelphia,  7,  1855, 

p.  440.    (Kauai.) 
O-O,  A-A 
Acridocercus  braccatus  Cassin  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  99; 

Perkins,  Fauna  Hawaiiensis  (Aves),  p.  445. 

Range.     Forests  of  Kauai.   Now  rare. 

Moho  nobilis  apicalis  Gould 

Moho  apicalis  Gould,  Proceedings  Zool.  Soc.  London,  1860,  p.  380.    (Owhyie 

in  error  =  Oahu.) 
Oahu  0-0 
Acridocercus  apicalis  Gould  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  103; 

Perkins,  Fauna  Hawaiiensis  (Aves),  p.  445. 


138  bulletin:  museum  of  comparative  zoology 

Range.  Formerly  forests  of  Oahu,  now  extinct.  Last  specimen 
collected  by  Herr  Deppe  in  1837. 

Moho  nobilis  bishopi  (Rothschild) 

Acrulocercus  bishopi  Rothschild,  Bulletin  British  Ornith.  Club,  1,  1893,  p.  xli- 

(Molokai.) 
Bishop's  0-0 
Acrulocercus  bishopi  Rothschild  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  Ill ; 

Perkins,  Fauna  Hawaiiensis  (Aves),  p.  445. 

Range.     Forests  of  Molokai.   Now  rare. 

Moho  nobilis  nobilis  (Merrem) 

Gracula  nobilis  Merrem,  Avium  rar  .  .  .  Icones  .  .  .,  1,  1786,  p.  7.   (Hawaii.) 

(fide  Davies-Sherborne) 
0-0  ' 
Acrulocercus  nobilis  Merrem  in  Wilson  and  Evans,  Aves  Hawaiiensis,  p.  105; 

Perkins,  Fauna  Hawaiiensis  (Aves),  p.  445. 

Range.     Formerly  forests  of  Hawaii.  Now  probably  extinct. 

Genus  Chaetoptila 

Chaetoptila  Sclater,  Ibis,  1871,  p.  358.  Type,  by  original  designation,  Entorniza 
?  angustipluma  Peale. 

Chaetoptila  angustipluma  (Peale) 

Entorniza  angustipluma  Peale,  U.  S.  Exploring  Expedition,  8,  1848,  p.  147. 

(Hawaii.) 
Kioea  (?) 

Range.  Formerly  mountain  forests  of  Hawaii.  Now  extinct. 
Recorded  from  "between  the  lower  Volcano  House  and  the  crater  of 
Kilauea"  where  Mills  collected  his  specimens  (fide  Rothschild  who 
quotes  Palmer  in  Birds  Laysan  Island,  etc.,  p.  216). 

"District  of  Hilo  near  Olaa"  (fide  Wilson  and  Evans,  Aves  Hawaii- 
ensis, p.  114). 

.  .  .  "supposed  to  have  come  from  Olaa,  it  is  very  doubtful  whether 
the  Chaetoptila  was  found  there.  It  is  much  more  probable  that  it 
was  confined  to  the  high  plateau  between  the  mountains  and  the 
upper  edges  of  the  forest  bordering  this,  where  it  was  observed  by 
Pickering  and  Peale  of  the  U.  S.  Exploring  Expedition"  (Perkins, 
Fauna  Hawaiiensis  (Aves),  p.  445).  All  these  speculations  are  open 
to  question. 


BRYAN  AND  GREENWAY:  HAWAIIAN'  BIRDS  139 

Part  II.     HYPOTHETICAL  FORMS  AND  RECORDS 

Family  ARDEIDAE— Herons 

Demigretta  sacra  (Gmelin) 

Ardea  sacra  Gmelin,  Systema  Naturae,  1,  pt.  2,  1789,  p.  640. 

Range.     Asiatic  coasts  and  islands  of  the  Pacific. 
Recorded  in  the  Hawaiian  Group  from  Kahalui,  Maui  (a  single  sight 
record  by  Finsch). 


Family  LARIDAE— Gulls 

What  is  now  thought  to  be  a  hybrid  Larus  hyperboreus  Gunnerus  x 
Larus  argentatus  vegae  Palmen  is  recorded  as  Larus  nelsoni  Henshaw 
(Auk,  1,  1SS4,  p.  250)  by  Oberholser  (Auk,  35,  1918,  p.  349)  from 
Hilo,  Hawaii. 

The  specimen  is  said  to  be  a  young  female  and  is  in  the  U.  S.  National 
Museum,  Washington. 

Subfamily  DREPANIDINAE 

Paroreomyza  perkinsi  (Rothschild) 

Oreomyza  perkinsi  Rothschild,  Avifauna  Laysan  Island,  etc.,  pt.  3,  1900,  p. 

129.   (Puulehua,  Hawaii.) 
Perkins's  Creeper 

Range.  Known  from  a  single  specimen,  the  type,  now  in  the 
American  Museum  of  Natural  History,  New  York. 

This  specimen  is  intermediate  in  color  and  form  of  bill  and  is,  in 
our  opinion,  a  hybrid  Chlorodrepanis  virens  x  Paroreomyza  mana. 
Although  the  feeding  habits  of  these  two  birds  differ,  we  do  not  think 
that  this  would  prevent  occasional  breeding,  Perkins  to  the  contrary 
(Fauna  Hawaiiensis  (Aves),  p.  417). 

Genus  "Sassius"  Rothschild  and  Hartert 

Sassius  Rothschild  and  Hartert,  Bulletin  British  Ornith.  Club,  46,  1926,  p.  51. 
Type,  by  monotypy,  Sassius  simplex  Rothschild  and  Hartert. 

"Sassius  simplex"  Rothschild  and  Hartert 

Sassius  simplex  Rothschild  and  Hartert,  Bulletin  British  Ornith.  Club,  46 
1926,  p.  51.    (Sandwich  Islands.) 


140  bulletin:  museum  of  comparative  zoology 

This  curious  little  skin  was  discovered  by  Dr.  Hartert  in  the 
Naturhistorisches  Museum  in  Vienna.  He  thought  it  to  be  an  un- 
described  Drepanid  at  the  time,  but  since  then  other  ornithologists 
have  thought  it  to  be  more  probably  an  artefact  made  of  the  skins 
of  Sun  Birds  (Nectariniidae),  an  African  and  Asiatic  family  (see 
Meise,  Proceedings  8th  Internat.  Orn.  Congress,  1934,  p.  123). 

Genus  "Dysmorodrepanis" 

Dysmorodrepanis  Perkins,  Annals  and  Magazine  Nat.  Hist.  (9),  3,  1919,  p.  250. 
Type,  by  monotypy,  Dysmorodrepanis  munroi  Perkins. 

"Dysmorodrepanis  munroi"  Perkins 

Dysmorodrepanis  munroi  Perkins,  Annals  and   Magazine  Nat.  Hist.   (9),  3, 
1919,  p.  251.    (Kaiholena  Valley,  Lanai.) 

This  curious  bird  is,  in  our  opinion,  an  aberrant  specimen  of 
Psittirostra  psittacca  (see  Greenway,  Auk,  56,  1939,  p.  479).  The  type 
and  only  specimen  is  in  the  B.  P.  Bishop  Museum,  Honolulu. 


THE  GENERIC  ARRANGEMENT  OF  THE  DREPANIDINAE 

By  James  C.  Greenway,  Jr. 

The  inter-relationships  of  the  genera  of  Drepanidinae  have  by 
some  been  considered  solved  by  the  work  of  R.  C.  L.  Perkins  Fauna 
Hawaiiensis  (Aves)  whose  arrangement  Sharpe  followed  In  his  Hand- 
List.  Perkins'  arrangement,  however,  is  not  entirely  satisfactory, 
for  in  some  cases  he  has  used  characters  which  form  what  appear  to 
be,  even  on  the  evidence  he  himself  uses,  rather  unnatural  groups. 

There  are  no  morphological  characters  to  prove  that  these  genera 
are  properly  placed  in  a  single  subfamily,  and  Perkins'  and  Gadow's 
works,  which  are  classical,  are  at  the  same  time  not  convincing,  since 
these  birds  have  no  single  character  in  common.  However,  the  ex- 
tremely divergent  genera  are  connected  by  intermediates,  and  the 
hypothesis  cannot  be  disproved.  Whether  the  group  sprang  from  a 
"Coerebine"  or  a  "Tanagrine"  stock,  or  both,  is  not  known.  It  is 
therefore  useless  to  speculate  about  the  specialization  of  forms. 

Within  the  subfamily  we  have  two  groups  which  may  be  called 
"Coerebine"  and  "Tanagrine",  simply  because  they  resemble  super- 
ficially these  groups.   The  first  have  tubular  tongues  with  well  devel- 


BRYAN  AND  GREENWAY:  HAWAIIAN  BIRDS  141 

oped  brush-like  tips.  This  group  includes  Viridonia,  Palmeria, 
Himatione,  Vestiaria,  Drepanis,  Hemignathus,  Heterorhynchus, 
Loxops  and  Chlorodrepanis.  At  this  point  there  is  a  sharp  demar- 
cation line,  for  the  tongues  of  the  following  genera  are  not  tubular, 
those  of  Paroreomyza  and  Ciridops  being  concave  and  with  a  modified 
brush  (from  this  point  of  view  intermediate).  Through  Psittirostra 
there  is  gradation  to  the  fleshy  tongues  of  Loxioides,  Rhodacanthis 
and  Telespiza,  which  are  the  markedly  "Tanagrine"  types.  In  spite 
of  the  sharp  difference  in  the  shape  of  the  tongue  there  is  no  other 
character  which  indicates  a  break  in  the  continuity  of  intermediate 
forms,  Chlorodrepanis  and  Paroreomyza  being  so  close  that,  were  it 
not  for  the  tongue,  they  would  be  considered  to  be  congeneric  under 
present  concepts  of  generic  limits.  Loxops,  too,  is  intermediate, 
having  the  bill  and  general  form  reminiscent  of  a  goldfinch  but  with 
a  typically  Drepanine  tongue  From  two  points  of  view  this,  is  rather 
an  aberrant  genus,  for  the  shape  of  bill  and  tongue  do  not  conform 
as  in  other  genera,  and  the  bill  with  its  "loxian  twist"  is  of  course 
very  curious.  According  to  Perkins,  it  is  a  convergent  parallelism  of 
both  habit  and  form. 

Perkins'  arrangement  groups  Drepanis,  Vestiaria,  Ciridops,  Palmeria 
and  Himatione  together,  and  this  assemblage  he  characterizes  as 
follows:  "The  genera  of  the  first  group  are  characterized  by  the 
truncate  apices  of  the  primaries,  except  in  the  anomalous  Palmeria, 
and  by  the  plumage  of  the  young  which  is  always  partly  black  or  of 
a  dull  colour."  This  is  an  unnatural  grouping  and  cannot  be  accepted. 
A  careful  examination  of  these  forms  reveals  the  fact  that  only 
Vestiaria  and  Himatione  have  truncated  primaries.  Furthermore, 
adults  of  Telespiza  ultima  are  melanistic  and  the  melanism  in  cantons 
is  quite  as  impressive  as  in  Vestiaria,  Perkins  to  the  contrary.  Melan- 
ism, in  any  event,  is  a  character  of  doubtful  value,  such  widely  diver- 
gent groups  as  Coereba  and  Charmosyna  being  classic  examples  of 
melanistic  mutation. 

Although  no  linear  arrangement  is  satisfying,  it  would  appear  to 
be  much  more  natural  to  place  the  tubular  ("Coerebine")  and  the 
more  fleshy  tongued  ("Tanagrine")  Drepanidinae  at  opposite  ends  of 
the  list,  with  the  intermediate  genera,  Chlorodrepanis,  Loxia,  Parore- 
omyza and  Ciridops  between  them.  Hemignathus  is  distinctly 
"Coerebine"  and  should  be  placed  close  to  Drepanis.  Ciridops  is  of 
course  a  very  curious  bird.  The  tongue  is,  however,  intermediate, 
and  even  though  the  color  pattern  (black  wings  and  tail)  resembles 
Vestiaria  slightly,  as  Perkins  remarks,  it  would  probably  be  better 


142  bulletin:  museum  of  comparative  zoology 

placed  as  an  intermediate  rather  than  with  the  typical  Drepanidinae. 
Ideally,  Chlorodrepanis  might  head  the  list,  leading  us  from  the 
Coerebinae  into  the  Drepanidinae. 

As  in  the  "Coerebine"  forms,  so  in  the  "Tanagrine"  forms  we  are 
gradually  led  from  a  tongue  which  in  Pseudonestor  approaches  a 
tubular  form  through  Psittirostra,  in  which  the  Drepanine  form  is 
still  observable,  to  Loxioides,  Rhodacanthis,  Telespyza  and  Chloridops. 
I  have  not  been  able  to  find  any  information  about  the  tongues  of  the 
last  two  genera,  but  Gadow  places  them  (p.  246)  near  Rhodacanthis 
which,  he  remarks,  has  the  "most  compact"  tongue  of  all.  There  is 
no  doubt  about  the  fact  (which  Gadow  points  out)  that  the  narinal 
structure  follows,  in  effect,  a  parallel  course. 

It  may  well  be  argued  that  the  last  four  genera  should  be  lumped 
in  a  single  genus,  as  Rothschild  suggests.  Loxioides  is  the  oldest  name. 
Lack  of  information  about  the  tongues  of  these  genera  deters  me 
from  this  course. 

No  new  facts  are  presented  here,  the  anatomical  information  having 
been  presented  by  Gadow  (in  Wilson  and  Evans,  Aves  Hawaiiensis), 
Rothschild  and  Perkins.  I  have  examined  tongues  of  Paroreomyza, 
Chlorodrepanis,  Himatione  and  Psittirostra.  The  interpretation  of 
evidence  collected  by  Perkins,  since  accepted,  seems  to  be  faulty, 
and  a,  to  me,  more  natural  linear  arrangement,  suggested  by  Gadow, 
follows : 

Viridonia 

Palmeria 

Himatione 

Vestiaria 

Drepanis 

Hemignathus 

Chlorodrepanis 

Loxops 

Paroreomyza 

Ciridops 

Pseudonestor 

Psittirostra 

Loxioides 

Rhodacanthis 

Telespyza 

Chloridops 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT   HARVARD    COLLEGE 

Vol.  XCIV,  No.  3 


OBSERVATIONS  ON  CHINESE  GOMPHINE 
DRAGONELIES 


By  James  G.  Needham 


CAMBRIDGE,  MASS.,  U.S.A. 

PRINTED    FOR    THE    MUSEUM 

June,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2 
have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT    HARVARD    COLLEGE 

Vol.  XCIV,  No.  3 


OBSERVATIONS  ON  CHINESE  GOMPHINE 
DRAGONFLIES 


By  James  G.  Needham 


CAMBRIDGE,  MASS.,  U.S.A. 

PRINTED    FOR    THE    MUSEUM 

June,  1944 


No.  4  —  Observations  on  Chinese  Gomphine  Dragonflies 
By  James  G.  Needham 

Chinese  dragonflies  first  caught  my  attention  in  1927,  when  I 
received  an  unanticipated  invitation  to  spend  a  year  in  China  visiting 
and  conferring  with  departments  of  Biology  in  the  Universities  of  that 
country  under  the  auspicies  of  the  China  Foundation  for  the  Promo- 
tion of  Education  and  Culture.  I  looked  about  for  aids  to  the  study  of 
the  local  dragonflies  and  found  there  were  none.  There  were  only  bare 
descriptions  of  the  adults  of  many  species,  printed  in  half  a  dozen 
languages,  and  well  scattered  through  the  zoological  literature  of  the 
world.   Nothing  was  known  of  the  immature  stages. 

Since  I  was  invited  to  lend  what  aid  I  might  to  the  study  of  biology 
in  China,  I  conceived  the  idea  that  by  supplying  a  manual  for  the 
study  of  the  one  group  of  insects  with  which  I  already  had  some  prac- 
tical acquaintance  at  home,  I  might  help  Chinese  students  in  the  study 
of  their  local  fauna.  Indeed,  I  was  quite  sure  that  such  aid  might  be 
more  real  and  the  results  more  lasting,  than  any  that  might  come  from 
merely  lecturing  to  them.  So  I  began  at  once  gathering  together  the 
materials  for  a  manual,  going  in  the  field  to  collect  dragonflies  as  I  had 
opportunity,  and  enlisting  the  aid  of  local  collectors  wherever  possible. 

I  started  with  no  collection  at  all,  and  with  almost  no  personal  ac- 
quaintance with  the  Oriental  Odonate  fauna.  The  literature  of  the 
group  was  largely  lacking  in  that  country.  Wherefore,  such  time  as 
could  be  taken  from  teaching  and  conferences,  I  devoted  to  collecting 
and  studying  dragonflies. 

After  my  arrival  in  September  I  did  a  little  collecting  of  autumnal 
species  in  and  about  Peiping,  with  the  invaluable  aid  of  Dr.  Chen-fu 
Wu  of  Yen  Ching  University,  and  Dr.  Chung-lo  Liu  of  Tsing  Hua 
University,  and  their  advanced  students.  They  guided  me  to  the  best 
collecting  places  in  some  of  the  most  beautiful  aquatic  situations  in 
North  China.  During  the  winter  I  was  able  to  work  on  my  collections 
in  the  private  laboratory  of  my  good  friend,  Dr.  N.  Gist  Gee,  who  was 
himself  a  lifelong  student  of  the  Chinese  fauna,  and  a  distinguished 
specialist  in  fresh-water  sponges.  In  the  spring  he  went  with  me  to  the 
Yang-tze  valley;  to  Soochow  and  to  Hang  Chow  Universities  where 
other  generous  collaborators  were  found.  I  spent  the  month  of  April 
in  Nanking;  that  was  my  real  harvest  season.  In  Nanking  I  had  the 
generous  assistance  of  Dr.  C.  Ping  and  a  number  of  his  research  stud- 
ents. There  I  had  considerable  time  for  collecting  adult  dragonflies, 
and  for  working  out  partial  life  histories.   In  May  I  returned  to  Peip- 


146  bulletin:  museum  of  comparative  zoology 

ing,  and  thereafter^war  conditions  prevented  further  field  work.  It 
was  with  profound  regret  that  I  had  to  leave  China  at  the  very  opening 
of  the  best  collecting  season.  Then  I  returned  home  to  America,  bring- 
ing all  the  collections,  made  and  borrowed.  Acknowledgment  is  made 
in  the  Manual  of  the  many  sources  of  the  borrowed  material. 

The  Manual  was  written  at  my  home  in  Ithaca,  in  such  intervals 
of  time  as  I  could  take  from  teaching  and  departmental  administra- 
tion. I  wrote  it  for  the  use  of  Chinese  students  in  the  study  of  their 
homeland  fauna.  I  sought  merely  to  provide  them  with  concise  de- 
scriptions, keys  and  tables  for  families,  genera  and  species;  and  in  all 
the  larger  genera,  where  species  are  difficult  of  identification,  to  pro- 
vide figures  of  the  genital  characters  that  are  the  ultimate  criteria  for 
species. 

In  matters  of  classification  I  considered  it  in  the  interest  of  students 
that  I  should  keep  to  the  older  and  simpler  family  groupings,  rather 
than  use  the  many  recently  introduced  and  still  untested  subdivisions 
of  the  families,  concerning  the  validity  of  which  the  specialists  are  not 
as  yet  in  entire  agreement.  I  wanted  to  provide  something  that  the 
college  students  could  use;  and  I  have  had  the  satisfaction  of  knowing 
that  they  have  used  the  Manual  successfully. 

In  only  one  group,  the  Gomphinae,  did  I  add  any  considerable  num- 
ber of  new  species.  Seventeen  of  these  were  in  the  large  and  hetero- 
geneous genus  Gomphus  of  the  older  authors.  Among  them  were  new 
heterogeneities  that  I  could  not  fit  to  some  of  the  subdivisions  of  that 
genus  that  had  been  recently  proposed.  I  contemplated  further  work 
upon  them  when  more  adequate  collections  and  more  time  should 
make  that  possible;  but  little  new  material  has  come  to  hand.  On 
completion  of  the  Manual  borrowed  specimens  were  returned  to  their 
owners ;  but  I  kept  duplicates  of  the  few  species  that  were  represented 
by  more  than  one  specimen;  and  I  took  occasion,  while  the  others  were 
in  my  hands,  to  make  photographs  of  the  wing  venation  of  most  of 
them.  And  now  that,  in  retirement,  I  have  time  for  more  adequate 
study,  I  have  only  this  scanty  material  available.  A  careful  restudy 
of  the  wing  venation  of  the  Chinese  Gomphines  is  hereinafter  at- 
tempted. 

The  venation  of  the  basal  half  of  the  hind  wing  appears  not  to  have 
been  thoroughly  explored;  for  in  it  there  are  structural  characters 
whose  taxonomic  value  has  been  quite  overlooked.  The  late  lamented 
E.  B.  Williamson  discovered  some  new  characters  in  these  parts,  and 
used  them  to  good  purpose  in  his  well  known  Burmese  paper  of  1907. 
There  are  yet  other  unused  characters  to  which  I  want  to  direct  at- 


NEEDIIAM:    CHINESE    GOMPHINE    DRAGONFLIES 


147 


tention  here.  To  them  I  will  apply  convenient  terms  for  use  in  the 
descriptions  that  are  to  follow.  The  new  terminology  will  be  merely 
supplemental  to  that  used  in  my  Manual  and  fully  illustrated  in 
Plate  I  of  that  volume. 

In  the  accompanying  figure  is  shown  a  diagram  based  on  a  careful 
drawing  of  the  base  of  the  hind  wing  of  Gomphus  campestris  Ndm., 
labelled  to  supply  names  for  the  parts  hereinafter  used.  The  principal 
longitudinal  veins,  Costa,  Subcosta,  and  Radius,  Media,  Cubitus  and 


Cu,  en, 

Fig.  1.    Base  of  the  hind  wing  of  Gomphus  (?  Burmagomphus*  campestris  Ndm. 


Anal,  are  labelled  in  the  figure  at  both  ends  of  each  vein,  with  appended 
numerals  designating  the  branches  of  some  of  them;  four  important 
cross  braces,  nodus  (n),  subnodus  (sn),  arculus  (ar),  and  anal  crossing 
(Ac)  and  three  other  parts,  middle  fork  (Mf),  anal  loop  (AL),  and 
membranule  (to),  illustrated  in  Plate  I  of  the  Manual,  are  repeated  in 
this  diagram. 

New  designations  here  introduced  are  as  follows:  intermedian  cross- 
f  ins  (i.  med.:  in  this  wing  there  is  but  one)  lie  in  the  space  between 
veins  M3  and  M4  and  beyond  the  arculus  to  the  middle  fork  (Mf). 

In  the  anal  area  of  both  wings  the  cells  that  lie  in  line  alongside  the 
anal  vein  from  base  to  gaff  (g)1  are  called  paranals.  In  the  fore  wing 
there  is  oftenest  but  a  single  row  of  them:  there  is  always  more  than 

1  The  gaff  is  the  fused  portion  of  veins  C'u2  and  Al. 


• 


148  bulletin:  museum  of  comparative  zoology 

one  row  in  the  hind  wing  and  the  cells  of  the  first  row  are  highly 
differentiated  and  are  of  great  systematic  importance.  In  our  diagram 
they  are  labelled  n,  o,  p,  and  AL.  Cells  n  and  o  are  constantly  present 
as  large  single  cells  in  all  Gomphinae  known  to  me,  with  the  single 
exception  of  Anormogomphus ;  cells  p  and  AL  may  be  divided  by  cross- 
veins.  An  anal  loop  (AL)  is  said  to  be  present  only  when  this  area  is 
definitely  bounded  by  a  strong  vein  in  the  rear  (with  that  boundary 
generally  much  stronger  than  shown  in  this  figure);  this  loop  is  often 
enlarged  and  divided  into  several  cells. 

For  the  cell  rows  running  in  the  opposite  direction,  from  front  to 
rear,  (more  especially  for  the  single  row  alongisde  vein  Al  extending 
from  the  hind  angle  of  the  triangle  to  the  wing  margin  and  numbered 
1,  2,  and  3  in  the  diagram)  I  use  Williamson's  name  post  anals. 

The  areas  into  which  the  broad  anal  field  of  the  hind  wing  is  divided 
by  the  three  branches  of  the  anal  vein  may  be  designated  as  the  first 
(x),  second  (y)  and  third  (z)  anal  interspaces.  Each  lies  behind  the 
branch  bearing  its  own  number  (principal  veins  being  numbered 
around  the  wing  margin  from  front  to  rear).  These  interspaces  differ 
in  breadth  and  slant,  and  in  number,  size  and  arrangement  of  the  cells 
composing  them.  Though  little  noticed  hitherto,  they  offer  excellent 
systematic  characters.  The  third  interspace  (z)  is  modified  into  the 
anal  triangle  of  the  male. 


GOMPHUS    s.lat. 

I  wish  now  to  make  a  further  analysis  of  the  species  that  I  lumped 
together  in  the  great  genus  Gomphus  in  the  Manual.  As  evidenced  by 
wing  venation,  these  species  fall  into  natural  groups  as  follows: 


Group  1.     G.  abdominalis  only 

For  it  I  propose  the  generic  name  GASTROGOMPHUS.  Its 
characters  are:  a  very  long,  thick  abdomen,  about  a  third  longer  than 
the  hind  wing;  anal  vein  3  arises  generally  after,  and  sometimes 
opposite  the  anal  crossing;  no  basal  subcostal  cross  vein,  and  no  cross 
veins  in  any  of  the  triangles;  first  and  fifth  antenodals  thickened;  a 
single  row  of  large  paranal  cells  in  the  fore  wing;  anal  triangle  of  the 
male  three  celled,  and  four  postanals  cells  in  the  hind  wing  (see 
Manual.  PI.  I,  fig.  4);  appendages  of  the  male  of  about  equal  length 
and  divergence  (Manual.  PI.  VI,  fig.  2). 


NEEDHAM:    CHINESE    GOMPHINE    DRAGONFLIES  149 

It  should  be  noted  also  that  in  the  one  known  species  there  is  a  very 
wide  differentiation  in  size  among  the  cells  of  three  wing  areas;  very 
large  before  the  level  of  the  areulus,  a  little  smaller  outward  to  a  line 
drawn  from  the  stigma  to  the  hind  angle  of  the  wing,  and  much  smaller 
thence  outward  to  the  margin. 

The  nymph  (Manual.  PI.  VII,  figs.  1  and  la)  differs  from  all  known 
related  forms  in  having  neither  dorsal  hooks  nor  lateral  spines;  in 
having  the  front  border  of  the  median  labial  lobe  doubly  produced 
(bilobed)  and  fringed  at  the  sides  of  a  bare  median  notch;  and  in 
having  the  strongly  incurving  terminal  third  of  the  lateral  lobe  very 
feebly  denticulate  on  its  concave  inner  margin. 

Group  2.     Xenogomphus,  gen.  now     Type  G.  agricola  Selys. 

Characters:  middle  fork  (Mf)  unsymmetrical,  askew  forward;  gaff 
as  long  as  or  usually  longer  than  the  inner  side  of  the  triangle;  inter- 
median  crossveins  3/1  in  fore  and  hind  wing  respectively;  anal  triangle 
of  the  male  hind  wing  usually  of  five  cells;  no  basal  subcostal  antenodal 
crossvein;  first  and  fifth  antenodals  thickened;  no  anal  loop,  but 
usually  two  complete  rows  of  cells  in  the  first  anal  interspace;  male 
caudal  appendages  of  equal  length  but  the  branches  of  the  inferior 
much  more  widely  outspread  (see  Manual.   PI.  VI,  figs.  6  and  6a). 

Here  belong  also  G.  succumbens  Ndm.  (Peking  Soc.  N.  H.  Bull.  5,  3, 
1930),  G.  citimus  Ndm.,  G.  lautus  Ndm.,  and  probably  also  (judging 
by  similarity  in  form  of  male  appendages)  G.  svrn-hcdini  Sjostedt  from 
Szechuan  and  G.  ekichibui  Fraser  and  G.  mclampus  Selys  from  Japan. 
This  is  the  only  group  of  species  in  the  Gomphus  series  of  Williamson 
that  has  the  middle  fork  (Mf)  unsymmetrical. 

The  nymph  (see  Manual  PI.  VII,  figs.  2  and  2a)  is  depressed  with 
strictly  lanceolate  abdomen  bearing  short  triangular  lateral  spines  on 
segments  7  to  9,  and  low  dorsal  hooks  on  segments  3  to  9,  the  latter 
very  small  at  the  front  and  regularly  increasing  in  size  to  rearward  to 
the  8th  segment.  The  middle  lobe  of  the  labium  is  prominent,  tri- 
angularly produced  with  a  pair  of  little  teeth  at  its  slightly  truncate 
tip,  and  fringes  of  marginal  hairs  at  either  side.  The  terminal  fourth 
of  the  lateral  lobe  beyond  the  base  of  the  strong  movable  hook  is 
roundly  incurved  to  meet  the  denticulate  inner  border,  without 
forming  a  distinct  terminal  hook. 

The  type  species  of  the  two  preceding  genera  are  both  pond  species, 
common  in  central  China,  where  I  made  rearings  of  both  of  them 
repeatedly. 


150 


bulletin:  museum  of  comparative  zoology 


Group  3.   Eogomphus,  gen.  nov.   Type  Gomphus  neglectus  Ndm. 

Characters :  Triangles  of  both  fore  and  hind  wings  long  in  the  axis 
of  the  wing,  and  generally  four-sided  by  failure  of  the  long  sides  to 
meet  at  the  outermost  angle ;  both  usually  traversed  by  a  single  cross 
vein;  bridge  vein  shortened  distally,  the  distance  from  subnodus  to 
oblique  vein  being  about  a  third  of  the  distance  of  subnodus  from  the 
middle  fork;  gaff  nearly  as  long  as  the  inner  side  of  the  hind  wing 
triangle;  basal  subcostal  crossvein  present  in  fore  wing,  absent  in  the 
hind;  vein  A2  weak  and  angulated  so  as  to  be  almost  unrecognizable; 


Fig.  2.    The  wings  of  male  (basal  part)  and  female  Eogomphus  neglectus  Ndm. 


for  a  considerable  distance  from  the  wing  margin  the  paired  long  veins 
inclose  more  than  one  cell  row,  the  greatest  doubling  between  M3  and 
M4,  less  between  Oil  and  Cu2,  least  between  Rs  and  M2;  behind 
Cu2  in  the  fore  wing  are  two  or  three  cell  rows  traversed  by  ill  de- 
veloped accessory  branch-like  sectors;  and  middle  fork  (Mf)  sym- 
metrical. 

This  genus  is  perhaps  nearest  to  Davidius  of  the  Gomphus  series,  but 
it  differs  in  having  the  fore  wing  triangle  longer  and  not  angulated  on 
the  outer  side;  the  intermedian  crossveins  are  little  reduced,  being  5/2 
in  fore  and  hind  wings  respectively.  This  character  transgresses  the 
lines  heretofore  drawn  between  the  Gomphine  and  Epigomphine 
series,  as  does  also  the  general  aspect  of  the  rather  elongate  wings. 

The  nymph  is  unknown.  The  nymph  figured  in  the  Manual  on 
Plate  VII,  figures  3  and  3a,  referred  to  on  page  67  as  possibly  belonging 
to  this  species,  is  the  nymph  of  Merogomphus. 


NEEDHAM:    CHINESE    GOMPHINE    DRAGONFLEIS  151 

The  three  teneral  specimens  representing  this  most  unusual  Gom- 
phine  came  to  me  just  before  the  manuscript  of  the  Manual  left  my 
hands.      One  male  was  retained  for  the  Cornell  University  Collection. 

After  a  reexamination  of  it,  I  may  here  add  another  note  of  de- 
scription. 

The  face  and  top  of  the  head  and  dorsum  of  the  first  abdominal 
segment  are  densely  hairy.  The  dorsum  of  the  bilobed  occiput  is 
thinly  clad  with  short  hair  that  is  parted  in  the  middle  and  outspread 
flat  both  ways  therefrom.  The  spines  on  femora  and  tibiae  are  very 
numerous,  short  and  in  uneven  alignment  in  the  outer  row  on  the  hind 
femur,  with  the  last  one  in  the  row  but  little  longer  than  the  others; 
those  of  the  middle  femur  are  more  than  twice  as  long,  equally  num- 
erous and  they  form  an  even  regular  row. 

The  superior  caudal  appendages  are  widely  divergent  from  the  base, 
tapering  and  convergent  only  toward  their  tips.  Each  ends  in  a  small 
black  tooth.  Below  the  base  is  a  large  inferior  branch  that  ends  in  a 
blunt  black  tooth.  The  inferior  appendage  is  little  more  than  half  as 
long  as  the  superiors  and  its  tips  have  less  than  half  their  spread.  It 
is  quadrangular,  with  a  straight  hind  margin  from  the  angles  of  which 
arise  two  stout  branches  that  project  straight  to  rearward.  Each  ends 
in  a  blunt  black  upturned  tooth.  The  genitalia  of  the  second  segment 
are  rather  prominent.  The  anterior  hamule  somewhat  resembles  the 
cheliped  of  a  craw-fish  with  both  tips  bluntly  rounded,  the  anterior  tip 
slightly  longer  and  inflexed  around  the  other.  The  posterior  hamule  is 
perhaps  twice  as  long  as  the  anterior.  There  is  a  bulbous  enlargement 
of  its  upper  third,  bearing  on  its  inner  side  a  cluster  of  about  a  dozen 
small  black  denticles;  then  suddenly  tapering  to  a  claw-like  incurved 
tip.  The  peduncle  ("vesicle")  of  the  penis  slopes  down  to  rearward, 
and  is  deeply  cleft  on  the  anterior  side  for  the  reception  of  the  greatly 
expanded  penis  tip.  It  is  nearly  bare  except  for  the  edges  of  this  cleft 
and  the  hood-like  inner  side  of  it.  The  second  joint  is  clavate  toward 
its  tip,  and  lacks  an  apical  spine.  The  reflexed  third  joint  bears  a  re- 
markable enlargement  at  its  tip;  a  deeply  cupped  expansion  that 
carries  a  suggestion  of  likeness  to  an  irregular  flower.  Above,  the  rim 
of  the  cup  is  deeply  emarginate,  and  within  it  arise  two  petal-like  lobes. 
Far  out  from  its  center  projects  a  bifid  stigma-like  process  ending  in 
a  pair  of  blunt,  recurved,  flabellate  tips,  below  which  projects  a  spine 
of  half  their  length. 

The  apical  carina  of  the  tenth  abdominal  segment  is  produced  to 
rearward  in  a  low  bare  triangular  prominence,  on  either  side  of  which 
is  the  usual  line  of  black  denticles. 


152  bulletin:  museum  of  comparative  zoology 

Group  4.     Merogomphus  Martin.  Type  M.  paviei  Martin 

A  more  careful  examination  of  the  single  known  female  specimen  of 
Gomphus  torpens  Ndm.  of  the  Manual  leads  me  to  conclude  that  it 
should  have  been  placed  in  this  genus  and  associated  with  Merogomphus 
vandykei  Ndm.,  for  it  has  the  following  characters :  vein  A3  arises  op- 
posite the  anal  crossing  (Ac),  and  not  beyond  it;  basal  subcostal  cross 
vein  (an)  present  in  the  fore  wing;  intermedial!  crossveins  3/1  in  fore 
and  hind  wing  respectively,  middle  fork  (Mf)  symmetrical;  gaff  more 
than  half  as  long  as  the  inner  side  of  the  triangle ;  two  rows  of  paranal 
cells  in  the  fore  wing;  first  anal  interspace  (x)  wider  than  the  second 
(y),  and  no  anal  loop.  It  also  has  a  half  a  dozen  very  large  spines  in  the 
outer  row  on  the  hind  femur. 

A  nymph  of  this  genus  was  figured  in  the  Manual  without  name 
(Plate  VII,  figs.  3  and  3a)  and  described  on  pages  66  and  67.  It  is 
probably  the  nymph  of  M.  vandykei,  as  determined  by  a  recent  study 
of  the  venation  of  its  crumpled  wings.1 

Group  5.  Mesogomphus  Foerster.  Type  Gomphus  cognatus  Rambur 

The  Gomphus  brevipennis  Ndm.  of  the  Manual  belongs  here,  as 
evidenced  by  the  following  characters:  vein. A3  arises  just  before  or 
opposite  the  anal  crossing  (.4c);  two  rows  of  paranal  cells  in  the  fore, 
wing;  intermedian  crossveins  2/1  in  fore  and  hind  wing  respectively; 
first  and  fifth  antenodals  thickened;  middle  fork  (Mf)  symmetrical; 
arculus  unusually  close  to  the  triangle  of  both  fore  and  hind  wings  and 
the  front  side  of  the  subtriangle  much  shorter  than  the  inner  side  of 
the  triangle;  four  postanal  cells  in  the  hind  wing;  four  or  five  cells  in 
the  male  anal  triangle;  no  anal  loop,  and  the  first  anal  interspace 
wider  than  the  second. 

The  nymph  has  been  described  and  figured  for  one  species  of  this 
genus,  M.  balneorum  by  Needham  and  Gyger  (Philippine  Jour.  Sci. 
63,  33,P1.X,  figs.  125  and  126)  and  for  two  others,  M.  lineatus  and  M. 
reinwardti  by  Lieftinck  (Tijd.v.Entomol.  77,21,  1934). 

Group  6.     Burmagomphus  Williamson.  Type  B.  williamsoni 

Fraser 

The  Gomphus  dolus  Ndm.  of  the  Manual  belongs  here.  In  venation 
it  is  very  close  indeed  to  the  type  species  as  figured  by  Williamson 

1  The  method  used  was  that  described  by  Dr.  May  K.  Gyger  in  Entomological  News,  50, 
p.  21,  1939. 


NEEDHAM:    CHINESE   GOMPHINE   DRAGONFLIES  153 

(Proc.  U.  S.  Nat.  Mux.  33,  298,  fig.  27,  1907).  The  combination  of 
venational  characters  by  which  this  genus  has  been  set  apart  is  as 
follows:  In  the  fore  wing,  a  single  row  of  paranal  cells;  a  small  triangle 
slightly  angulated  near  the  middle  of  its  outer  side;  a  long  close 
parallel  of  veins  M4  and  (Ail  beyond  it  with  but  two  intervening 
rows  of  cells  out  to  the  level  of  the  oblique  vein,  with  a  rather  sudden 
widening  thereafter;  and  in  the  hind  wing  but  three  postanal  cells. 

The  nymph  of  this  genus  has  been  mentioned,  and  given  an  un- 
intelligible two-line  description  by  Fraser  in  Fauna  of  British  India: 
Odonata  2,  212,  1934. 

Another  small  species  that  would  appear  to  belong  near  to  Burma- 
gomphus  is  the  one  a  portion  of  whose  hind  wing  is  shown  in  the 
accompanying  figure  of  Gomphus  campestris  Ndm.  of  the  Manual. 
(See  figure  1.)  It  is  of  small  size  (hind  wing  25  mm),  with  slightly 
angulated  outer  side  to  the  hind  wing  triangle,  and  only  three  postanal 
cells.  The  single-celled  anal  loop  is  quite  like  that  of  B.  icilliamsoni. 
There  are  these  discordant  characteristics:  there  is  a  basal  subcostal 
crossvein  in  both  fore  and  hind  wings;  one  or  two  cells  of  the  paranal 
row  in  the  fore  wing  are  divided,  and  the  double  row  of  cells  beyond  the 
triangle  does  not  extend  outward  beyond  the  level  of  the  nodus.1 
The  venation  as  a  whole  is  more  sparse,  there  being  but  11:7/8:7 
nodal  crossveins  in  fore  and  hind  wing  respectively  and  only  three 
crossveins  under  the  bridge. 

My  material  is  inadequate  for  determining  whether  these  differ- 
ences are  constant  enough  to  justify  generic  separation.  Since  this 
species  has  been  taken  on  the  campus  of  Yen  Ching  University,  it 
should  be  possible  for  some  one  there  to  obtain  additional  specimens 
including  also  its  immature  stages. 

Group  7.  Gomphus  s.  str.  Type  Libellula  vulgatissima  Linn 

The  remaining  species  appearing  under  this  generic  name  in  the 
Manual  may  be  allowed  to  remain  so  for  the  present.  They  show  a 
general  conformity  to  the  type,  but  with  numerous  small  divergencies 
which  I  shall  now  try  to  indicate  in  so  far  as  they  appear  in  the  vena- 
tion of  the  wings.  They  all  seem  to  have  the  following  characters  in 
common:  middle  fork  (Mf)  symmetrical;  vein  A3  arises  beyond  the 
anal  crossing  (Ac);  intermedian  crossveins  generally  2/1;  paranal 
cells  in  the  fore  wing  more  than  a  single  row,  some  cells  at  least  being 

1  This  disagreement  applies  also  to  Fraser's  figure    of  Burmagomphus    pvramidalis,  Faun. 
British  India  Odonata:  2,  212,  fig.  66,  1934. 


154  bulletin:  museum  of  comparative  zoology 

divided;  postanal  cells  four  to  seven;  anal  triangle  of  the  male  gener- 
ally three  celled;  and  no  basal  subcostal  cross  veins. 

Recalling  Burniagomphus  are  the  two  small  shortwinged  species 
G.  arvalis  Ndm.  and  G.  sowerbyi  Ndm.  These  have  a  well  defined  one 
celled  anal  loop  with  the  base  of  vein  Al  kinked  around  its  outer 
corner.  The  outer  side  of  the  nearly  equilateral  fore  wing  triangle  is 
slightly  angulated  in  the  middle.  The  arculus  is  between  the  first 
and  second  antenodal  cross  veins.  The  gaff  is  about  half  as  long  as 
the  inner  side  of  the  hind  wing  triangle.  Vein  M4  is  slightly  undulate, 
and  there  are  four  post  anal  cells.  This  latter  character  prevents  plac- 
ing them  in  Platygomphus,  as  does  also  the  well  developed  3-celled 
anal  triangle  in  the  male.  De  Selys  placed  a  question  mark  before  his 
Platygomphus  occultus  when  he  placed  it  in  that  genus.  It  belongs 
rather  with  the  above  named  pair.  The  three  might  possibly  be  made 
the  basis  of  a  new  genus;  but  until  Burmagomphus  and  Platygomphus 
are  better  defined,  and  until  Gomphus  campcstris  has  found  its  place, 
and  until  the  nymphs  of  all  of  them  are  made  known,  another  name 
would  only  add  to  the  confusion. 

Two  somewhat  larger  species  of  the  Manual,  G.  intinctus  and  G. 
collar  is,  are  like  the  three  preceding  in  most  characters,  but  the 
arculus  is  nearer  the  second  antenodal  crossvein,  and  intinctus  has 
five  postanal  cells.  In  all  five  the  gaff  is  about  half  as  long  as  the  inner 
side  of  the  triangle. 

A  peculiar  species  that  is  known  unfortunately  from  only  a  single 
female  specimen  is  G.  edax  Ndm.  The  triangles  are  both  elongated  in 
axis  of  the  wings,  the  outer  end  of  the  hind  wing  triangle  being  turned 
up  at  the  end  like  a  sled  runner  (see  the  next  figure) ;  there  are  seven 
postanal  cells ;  the  branches  of  the  anal  vein  are  aslant  outward. 

Another  peculiar  species,  described  later  by  me  (lAngnan  Sci.  Jour. 
10,  227,  1931)  from  a  single  female  specimen  taken  in  Hainan  is 
G.  hoffmani.  It  has  the  first  and  sixth  antenodals  thickened,  the 
arculus  at  or  beyond  the  second;  no  basal  subcostal  antenodal  cross- 
vein,  the  fore  wing  with  two  rows  of  paranal  cells,  no  large  cells  in  the 
anal  area  behind  the  first  paranal  row,  and  the  first  anal  interspace 
much  wider  than  the  second.  Added  to  this  array  of  differences  there 
is  a  peculiarly  elongated  three-celled  anal  loop,  around  the  outer  end 
of  which  vein  Al  makes  a  short  sharp  bend.  Also  the  hind  wing  is 
widest  at  the  nodus.  The  gaff  is  as  long  as  the  inner  side  of  the  tri- 
angle, which  latter  in  the  fore  wing  is  a  little  longer  than  the  front  side. 
Whether  this  wing  is  quite  normal  I  cannot  say. 

There  remain  six  large  species  (hind  wing  37-40  mm.)  that  conform 


NEEDHAM:    CHINESE    GOMPHINE   DRAGONFLIES  155 

more  closely  (still  none  too  well)  to  the  type  of  the  genus.  One, 
G.  cuneatus  Xdm.  of  the  Manual  appears  to  be  distinguished  by  having 
the  veins  M3  and  M4  not  at  all  undulate,  strictly  parallel  and  regularly 
curved  and  inclosing  somewhat  larger  cells  than  in  the  other  five 
species.  In  general  it  has  a  more  open  venation.  It  also  has  a  longer 
gaff,  almost  as  long  as  the  inner  side  of  the  triangle,  and  a  shorter 
kink  in  the  base  of  the  anal  vein  at  the  outer  angle  of  the  one  celled 
anal  loop.  There  is  in  the  type  specimen  but  one  extra  cell  in  the 
otherwise  single  paranal  row  of  the  fore  wings. 

Of  the  five  remaining  species,  G.  endicotti  appears  to  be  separable  by 
reason  of  the  shortness  of  the  front  side  of  the  fore  wing  triangle — not 
longer  than  the  inner  side  of  the  same;  and  G.  flavicomis  (Peking  Soc. 
Nat.  Hist.  Bull.  1,  2,  1930)  by  having  its  anal  crossing  close  to  the 
inner  end  of  the  subtriangle  in  the  hind  wing — less  than  half  its  own 
length  therefrom. 

Finally  G.  amicus  is  separable  from  the  remaining  two  by  the 
shortness  of  its  gaff — less  than  half  the  length  of  the  inner  side  of  the 
front  wing  triangle;  and  these  last  two  may  be  separated  by  the  posi- 
tion of  the  arculus  in  relation  to  antenodal  crossveins:  it  is  midway 
between  the  first  and  second  in  G.  clathratus,  and  at  or  very  close  to 
the  second  in  G.  septimus. 


THE  FRAMEWORK  OF  THE  WING  ABOUT  THE 

TRIANGLES 

In  the  preceding  pages  I  have  been  pointing  out  the  best  single  vena- 
tional  characters  that  I  have  been  able  to  find  for  distinguishing  each 
of  the  species  listed  in  my  Manual  under  GOMPHLS  (all  of  them  ex- 
cept G.  sornnolens,  of  which  I  now  have  neither  wings  nor  photographs 
of  venation  available).  I  now  wish  to  present  in  the  form  of  a  table 
some  correlations  of  characters  especially  to  show  the  relations  of  the 
parts  of  the  strong  framework  of  the  wing  that  are  in  or  around  the 
triangles. 

As  a  standard  of  comparison  I  take  the  part  marked  a,  which  forms 
the  one  common  side  of  triangle  and  subtriangle,  and  which  is  formed 
in  development  about  the  posteriorly  deflected  portion  of  the  main 
Cubital  trachea.  Two  additional  reasons  for  its  selection  are  (1)  its 
central  location,  its  ends  being  in  contact  with  all  the  other  parts 
compared;  and  (2)  its  relative  constancy  in  length.  A  little  comparison 


156 


bulletin:  museum  of  comparative  zoology 


showed  all  the  other  parts  to  be  more  variable.  Next  in  constancy  was 
the  part  marked  e,  which  is  formed  about  the  main  Anal  trachea. 

The  part  a  was  given  an  assigned  value  of  10,  and  the  length  of  all 
the  other  parts  were  estimated  in  tenths  of  it.  That  is  the  meaning  of 
the  numerals  in  the  central  columns  of  the  table.    These  values  are 


Fig.  3.    The  parts  about  the  triangle  (T)  in  the  wing  hind  of  Gomphus  edax  Ndm.  to  illustrate 
the  terms  used  in  the  following  table. 


merely  estimates  made  under  inspection  with  a  lens,  without  careful 
measurements:  wherefore  allowance  (possibly  up  to  10%)  may  have 
to  be  made  for  errors  of  judgment,  and  additional  allowance  for  varia- 
tion in  individual  specimens. 

The  accompanying  enlarged  diagram,  based  on  a  drawing  of  the 
wing  of  Gomphus  edax  Ndm.  will  be  useful  for  comparison. 

In  general  it  may  be  said  concerning  venational  characters  that  the 
conjunctions  and  proportions  and  directions  of  the  component  parts 
of  the  strong  framework  of  the  wing,  and  the  layout  of  the  spaces  be- 
tween principal  veins  and  their  branches  offer  far  more  dependable 
taxonomic  characters  than  are  to  be  found  in  the  number  of  interven- 
ing crossveins. 


NEEDHAM:    CHINESE    GOMPHIXE   DRAGOXFLU SS 


157 


Venational  Characters  in  20  Species  of  Gomphus 


Species 

Hind 
wing 

I n  ter- 
med. CVS. 

f/h  tv. 

a 

Length  of 

parts  about 

hind  W.  triangle 

b      e      d      e    f 

9 

extra 
paran. 

lis 

post 
a  rials      A3sAc 

A.  loop 
present 

abdominalis 

35 

2/1 

10 

17  18     9 

11  5 

5 

0 

4 

opp 

yes 

agricola 

25 

3/1 

10 

16  18  10 

11  6 

12 

0 

4 

out 

no 

amicus 

40 

2/1 

10 

16  18  11 

12  8 

4 

3 

4 

in 

yes 

arvalis 

29 

2/1 

10 

18  20  12 

12  7 

4 

2 

4 

in 

yes 

campestris 

clathratus 

collaris 

26 
38 

31 

2/1 
3/1 
2/1 

10 
10 
10 

12  13  11 
17  19  10 
15  16  10 

12  5 
11  7 

11  7 

5 
5 

4 

1 
2 

2 

3  • 

4 
4 

n 
n 

n 

yes 
yes 
yes 

cuneatus 

38 

2/1 

10 

15  16  10 

11  7 

7 

1 

5 

in 

yes 

dolus 

23 

2/1 

10 

9  11     9 

11  4 

6 

0 

3 

m 

no 

edax 
endicotti 

35 
34 

3/1 

2/1 

10 
10 

18  20  11 
17  19  10 

12  6 
11  7 

8 
9 

2 
3 

5         i 
5         i 

n 

n 

yes 
yes 

flavicornis 

37 

2/1 

10 

17  19  10 

11  5 

4 

2 

5         ] 

n 

yes 

gideon 
hoffmanni 

36 
34 

4/2 
2/1 

10 
10 

16  17  10 
15  16  10 

11  6 

11  7 

5 
9 

3 
4 

4 
5 

n 
n 

yes 
no 

intinctus 

31 

2/1 

10 

14  15  10 

11   7 

4 

4 

5 

in 

yes 

neglectus 

36 

5/2 

10 

19  20  10 

11  5 

7 

0 

5 

out 

no 

occultus 

30 

2/1 

10 

14  15     9 

10  5 

7 

1 

4 

m 

yes 

septimus 
sowerbyi 

•  40 
29 

2/1 
2/1 

10 
10 

13  15     9 

14  15     9 

11  6 
11  6 

7 
4 

3 
2 

5 

4 

in 
in 

yes 
yes 

torpens 

30 

3/1 

10 

15  16     9 

11  4 

10 

2 

4       i 

n 

no 

Column  1.    Length  of  hind  wing  in  millimeters. 

Column  2.    Number  of  crossveins  joining  the  sectors  of  the  arculus  between  the  arculus 
and  the  middle  fork  in  fore  and  hind  wing. 

Column  3.    Relative  lengths  of  the  parts  about  the  triangle  of  the  hind  wing  in  terms  of 
tenths  of  the  length  of  the  inner  side  of  the  triangle. 

Column  4.    Number  of  extra  paranal  cells  in  the  front  wing  (more  than  the  single  row  always 
present). 

Column  5.    Number  of  cells  in  the  postanal  row  on  the  proximal  side  of  vein  1st  A  between 
the  triangle  and  the  hind  margin  of  the  wing. 

Column  6.    Position  of  origin  of  vein  3d  A:  in,  proximal  to  it;  opp,  opposite  it;  out,  distal 
to  it. 

Column  7.    Anal  loop  of  hind  wing. 


158 


bulletin:  museum  of  comparative  zoology 


Leptogomphus  unicornis  Ndm. 

A  study  of  the  wings  of  the  single  known  specimen  of  this  species 
shows  it  to  have  been  misplaced  in  the  Manual  in  the  genus  Davidius. 
It  has  more  in  common  with  Leptogomphus,  including  (1)  the  form  of 
the  wings;  (2)  the  lack  of  brace  vein  to  the  stigma;  (3)  the  trigonal 
interspace  regularly  widening  outward  to  the  wing  margin;  (4)  the 
small  triangles ;  (5)  the  narrow  fore  wing  subtriangle ;  (6)  the  little  ex- 
panded anal  area  of  the  wing;  and  (7)  the  3rd  anal  interspace  (z) 
longer  in  the  axis  of  the  wing  than  wide.  All  these  I  regard  as  primitive 
characters,  of  relative  fixity. 


Fig.  4.    Wings  of  ? Leptogomphus  unicornis  Ndm. 


It  seems  to  differ  from  Leptogomphus  as  represented  by  its  type 
species,  L.  sauteri  Selys,  in  having  but  a  single  row  of  very  large 
paranal  cells  in  the  fore  wing,  in  having  no  basal  subcostal  antenodal 
crossvein,  and  in  having  crossveins  in  all  the  triangles  and  in  the 
supratriangular  space  of  both  the  wings.  I  present  a  figure  of  the  vena- 
tion to  call  attention  to  these  discrepancies.  At  first  glance  they  seemed 
to  me  to  be  so  great  as  to  call  for  generic  separation;  but  on  further 
experience  with  the  Epigomphus  alliance,  I  think  they  are  very  un- 
reliable variants,  having  only  specific  value. 


NEEDHAM:    CHINESE    GOMPHINE   DRAGONFLIES  159 


TWO  NEW  SPECIES  AND  SOME  NEW  RECORDS 

Since  the  publication  of  the  Manual  and  of  the  Additions  and  Cor- 
rections (in  the  Peking  Soc.  of  Nat.  Hist.  5,  1-10,  1930)  only  one  small 
collection  of  dragonflies  has  come  to  me  from  China.  It  was  from  Dr. 
Ting-wei  Lew.  It  contained  two  new  species  of  Gomphines  and  es- 
tablished a  few  new  records. 


Gomphurus  gideox  spec.  nov. 

Length  63  mm.;  abdomen  45;  hind  wing  38. 

This  is  a  blackish  species  with  spatulately  dilated  abdomen;  face 
black  with  an  oblong  stripe  of  yellow  covering  about  half  of  the  lab- 
rum.  A  similar  somewhat  larger  isolated  stripe  covers  the  top  of  the 
frons.   The  remainder  of  the  top  and  rear  of  the  head  is  black. 

The  synthorax  is  black  in  front  with  a  pair  of  isolated  dorsal  stripes 
that  are  divergent  downward,  not  reaching  the  divided  cross  stripe  on 
the  collar.  An  antehumeral  stripe  of  yellow  is  represented  by  a  very 
small  spot  high  up  near  the  crest,  and  a  thin  faint  streak  low  down  on 
the  side,  its  lower  end.  Behind  the  broad  black  humeral  band  the 
sides  are  yellow  with  a  black  line  on  the  third  lateral  suture,  that  con- 
nects with  the  back  subalar  carina  above,  and  below  runs  down  behind 
the  coxa  of  the  middle  legs.  A  vestige  of  a  middle  stripe  is  present  in 
front  of  the  spiracle.  Legs  black  beyond  the  short  bicolored  basal 
segments.  Wings  hyaline,  with  a  faint  tinge  of  yellow  in  the  mem- 
brane. Ante-  and  postnodal  crossveins  are  16:12  and  10/12  in  fore 
and  hind  wing  respectively. 

The  abdomen  is  very  moderately  enlarged  on  the  two  basal  segments 
narrowly  cylindric  on  segments  3  to  7,  and  spatulate  on  7  to  10,  with 
widely  flaring  lateral  expansion  of  the  margins  of  7,  8  and  9.  The 
dorsum  of  1  is  mainly  yellowT  and  a  narrow  lanceolate  spot  appears  on 
the  base  of  2.  The  sides  of  1  and  2  are  mainly  yellow,  and  also  the  base 
of  3.  Basal  yellowish  rings  on  3  to  7  become  narrower  to  rearward, 
with  only  fine  yellow  intersegmental  lines  across  the  apices  of  7,  8, 
and  9:  segment  10  and  appendages  wholly  black.  The  relative  length 
of  the  last  four  segments  middorsally  is  about  as  11;  9:  10:5;  and  ap- 
pendages, on  the  same  scale  as  7.  Diffuse  large  yellow  spots  cover 
about  half  of  the  sides  of  segments  8  and  9.  Caudal  appendages  are  as 


160  bulletin:  museum  of  comparative  zoology 

shown  in  figure.  The  posterior  hamules  of  the  male,  completely  hiding 
the  anterior  ones,  project  strongly  downward  even  beyond  the  level 
of  the  "vesicle",  and  taper  to  claw-like  sharp  tips  that  are  directed 
forward. 


Fig.  5.    The  abdomen  of  Gomphurus  gideon  sp.n.,  dorsal  view. 

The  female  is  very  similar  in  coloration,  with  the  yellow  areas  a  little 
more  extended,  especially  on  the  abdomen.  The  subgenital  plate  is 
divided  deeply  into  two  blunt  equilateral  triangles  that  extend  to 
rearward  across  about  a  fifth  of  the  9th  sternite.  The  supraanal  plate 
is  shining  black  above,  yellow  beneath. 

A  single  pair,  type  and  paratype,  collected  in  Chengtu,  Szechuan 
on  June  29th,  1929,  and  sent  me  by  Dr.  Lew.  They  are  now  in  the 
Cornell  University  Collection. 

Because  of  the  striking  dilatation  of  the  7th  and  8th  segments  of  the 
abdomen,  shown  in  the  figure  herewith,  and  general  accord  in  caudal 
appendages  and  in  venation,  I  have  placed  this  species  in  Gomphurus; 
a  genus  hitherto  known  only  from  North  America.  Another  Chinese 
species,  Gomphus  kryenbergi  Ris,  compared  by  its  describer  with  the 
American  Gomphus  scudderi,  doubtless  belongs  beside  it.  G.  gideon  has 
however  a  somewhat  more  copious  venation  than  the  American 
species,  with  four  cells  in  the  anal  triangle  in  both  right  and  left  hind 
wings  of  the  male:  intermedian  cross  veins  4/2  in  both  male  and  female, 
and  six  postanal  cells. 


NEEDHAM:    CHINESE    GOMPHINE   DRAGONFLIES  161 

Davidius  serenus  spec.  nov. 

Female;  length  41  mm.;  abdomen  31;  hind  wing  27. 

This  is  a  small  blackish  species  with  yellow  sides.  The  head  is  all 
black  except  the  outer  sides  of  the  mandibles  and  a  broad  transverse 
stripe  across  the  very  low  prominence  of  the  frons,  which  are  yellow. 
The  thorax  is  black  in  front  except  for  a  pair  of  opposed  7-marks  that 
just  meet  at  the  middle  of  the  collar.  The  stalks  of  the  7s  are  slightly 
tapered  upward  and  blunt  at  their  isolated  upper  ends. There  is  no 
antehumeral  yellow  stripe  at  all.  Behind  the  very  broad  black  humeral 
band  the  sides  are  mainly  yellow  with  a  narrow  black  stripe  on  the 
third  lateral  suture  that  is  connected  forward  with  the  humeral  above 
and  below.  The  black  of  the  ventral  side  extends  upward  at  the  middle 
suture  to  cover  the  spiracle.  The  legs  are  black.  The  long  slender  hind 
femora  are  sagged  downward  in  the  middle  and  beset  underneath  with 
more  than  a  score  of  slender  black  short  subequal  spines. 

Wings  hyaline.  Ante-  and  postnodals  13:12/9:11  in  fore  and  hind 
wing  respectively.  There  is  an  extra  cubito-anal  crossvein  in  the  fore 
wing,  and  there  is  a  single  row  of  cells  behind  the  anal  vein. 

Abdomen  mostly  black  with  a  diminishing  amount  of  yellow  on  the 
sides  of  segments  1  to  7;  segment  1  mostly  yellow  dorsally  and  2  with 
a  lanceolate  streak  of  the  same  color.  On  the  sides  of  segments  3  to  5 
the  yellow  is  broken  into  a  row  of  three  spots;  reduced  to  two  spots  on 
6,  and  to  a  single  spot  on  7:  8  to  10  black.    Appendages  yellow. 

The  subgenital  plate  of  the  female  is  oblong  flat,  slightly  tapering  to 
rearward,  with  a  deep  notch  at  the  tip,  and  about  four  fifths  as  long 
as  the  venter  of  the  9th  segment. 

This  species  is  nearly  allied  to  D.  trox  Ndm.,  but  differs  in  being 
smaller  in  size,  in  having  the  labium  all  black,  in  lacking  the  pale  spot 
that  is  a  vestige  of  the  antehumeral  stripe,  in  lacking  the  J-spot  at  the 
rear  of  the  side  of  the  thorax,  and  in  having  the  abdominal  segments 
wholly  black  and  the  appendages  yellow. 

There  is  a  single  specimen  collected  at  Ruling,  China  in  July  1933 
by  Dr.  Ting-wei  Lew,  and  now  in  the  Cornell  University  collection. 

Among  the  specimens  sent  me  by  Dr.  Lew  were  five  females  of 
Gomphus  septimus  Ndm.  The  male  was  described  in  the  Manual,  p.  61. 
The  female,  heretofore  unknown,  is  like  the  male  in  coloration,  with 
a  basal  yellow  halfring  on  abdominal  segment  7  more  conspicuous  than 
in  the  male.  The  relative  length  of  the  last  four  abdominal  segments 
is  as  15:12:10:7,  with  the  appendages  8,  on  the  same  scale.  The  promi- 
nent subgenital  plate  is  scoop-shaped  or  shaped  like  the  spout  on  a 


162  bulletin:  museum  of  comparative  zoology 

pitcher,  triangular,  black,  more  than  half  as  long  as  the  venter  of  the 
9th  segment,  and  directed  conspicuously  downward.  Among  the  rather 
stout  spines  on  the  hind  femur  are  four  to  six  stronger  than  the  others, 
but  intermixed  with  the  others,  and  none  of  them  is  as  long  as  the 
femur  is  thick. 

Two  of  the  specimens  came  from  Foochow  in  May;  one  from  Mt. 
Poliang  ding,  near  Ho-kiang  and  Ming-kiant  Fukein;  and  two  from 
Tu-ching,  Min-giang  in  Fukien. 

Dr.  Lew  sent  also  a  fine  pair  of  Mcgalogomphus  sommeri  Selys  from 
Kuling;  a  species  that  has  hitherto  been  without  definite  locality 
assignment  in  China. 


TWO  CORRECTIONS  FOR  THE  MANUAL  AND 
A  CONFIRMATION 

(1)  Agricnemis  amelia  Ndm.  (Manual,  p.  256)  is  a  synonym  of 
Ischnura  delicata  Hagen,  which  is  in  turn  considered  by  some  authori- 
ties to  be  a  synonym  of  Ischnura  aurora  Burmeister. 

(2)  Taolcstes  ncctans  Ndm.  (Manual,  p.  256)  is  correctly  described 
and  illustrated,  but  the  nymph  associated  with  it  belongs  elsewhere. 
The  female  type  specimen  was  presented  to  me  along  with  a  cast 
nymphal  skin,  supposedly  reared,  and  with  several  nymphs  that  had 
been  collected  at  the  same  spot.  I  uncritically  accepted  them  as  be- 
longing together.  The  nymphs  were  not  well  preserved;  but  in  a 
recent  examination  of  one  of  the  best  of  them  I  find  enough  venation 
still  remaining  in  its  wing  pads  to  show  that  the  antenodal  crossveins 
are  numerous.   That  is  sufficient  to  show  that  they  are  not  Taolcstes. 

The  nymph  is  structurally  very  similar  to  those  now  known  belong- 
ing to  the  genus  Euphaca.  Judging  by  its  size,  it  should  be  Euphaca 
opaca. 

A  word  about  the  placement  of  Taolcstes.  As  explained  at  the  be- 
ginning of  this  article,  I  followed  the  older  and  easier  system  that 
segregated  the  Lestinae  from  the  others  on  one  principal  character: 
middle  fork  (Mf)  nearer  the  arculus  than  the  nodus.  But  in  doing  so 
I  pointed  out  (p.  226)  the  nonconformity  of  Taolcstes  with  the  true 
Lestinae. 

Two  additional  species  have  since  been  described  from  China  by 
Erich  Schmidt  (Koitowia  10,  178-183,  1931)  as  species  of  Rhipidolcstes: 
R.  bidens  and  R.  truncatidens;  they  conform  much  more  closely  to 
Taolcstes,  not  only  in  having  the  middle  fork  nearer  the  arculus  than 


NEEDHAM:    CHINESE    GOMPHINE    DRAGONFLIES  163 

to  the  nodus,  but  also  in  many  other  points  of  venation,  and  in  the 
male  genitalia.  The  species  T.  nectans  differs  from  both  in  having  un- 
marked hyaline  wings,  much  more  open  venation,  and  a  shorter  and 
thicker  stigma. 

Concerning  the  nymph  of  Megalestes,  I  noted  on  page  229  of  the 
Manual  that  Laidlaw  had  described  a  nymph  that  he  referred  by  sup- 
position to  M.  major  (Ind.  Mus.  Record,  19,  185-187,  1920).  I  said  I 
was  not  convinced  that  Laidlaw's  nymph  belonged  to  Megalestes. 
My  remark  was  more  than  justified:  this,  notwithstanding  Leiftinck's 
oracular  pronouncement  (Treubia  17,  58-61,  1939)  in  support  of  Laid- 
law's supposition.  I  obtained  an  almost  identical,  certainly  congeneric 
nymph  of  Rhinagrion  philippinum  from  Luzon,  well  preserved,  and 
showing  so  complete  venation  in  its  wing  pads  as  to  leave  no  doubt  as 
to  its  identity.  It  is  described  and  figured  in  the  Philippine  Journal  of 
Science,  70,  266,  Plate  15,  figs.  206-213  and  215-216,  1939.  Laidlaw's 
nymph  is  Rhinagrion. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT   HARVARD   COLLEGE 

Vol.  XCIV,  No.  4 


P3AMMOCHARIDAE 

(Spider-Wasps) 

Notes  and  Descriptions 


By  Nathan  Banks 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED   FOR   THE  MUSEUM 

June,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


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Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2 
and  3  have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

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Bulletin  of  the  Museum  of  Comparative  Zoology 

AT   HARVARD   COLLEGE 

Vol.  XCIV,  No.  4 


PSAMMOCHARIDAE 

(Spider-Wasps) 

Notes  and  Descriptions 


By  Nathan  Banks 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED   FOR  THE  MUSEUM 

June,  1944 


No.  4  —  Psammocharidae1 

(Spider- Wasps) :  Notes  and  Descriptions 

By  Nathan  Banks 

In  continuation  of  previous  studies  on  the  fine  collection  of  Psam- 
mocharidae in  the  Museum  of  Comparative  Zoology  I  present  descrip- 
tions of  some  new  genera  and  species  and  notes  on  others.  Neotypes 
are  made  for  two  of  Say's  species  and  descriptions  of  varietal  forms. 
Some  of  these  varietal  forms  illustrate  the  faunal  divergence  of  the 
New  England  area,  which  I  have  previously  noted  in  a  Myrmeleonid, 
Hesperolcon  abdominalis  Say.  Descriptions  are  given  of  the  males  of 
three  West  Indian  species  previously  known  only  from  the  females. 
Keys  are  made  for  the  species  of  Dipogon  and  for  the  males  of  the 
species  of  Ageniella  known  to  me  from  the  Eastern  States.  A  compari- 
son of  our  large  black  Arizona  Pepsis  with  a  series  of  both  sexes  of  the 
related  Cuban  form  and  the  South  American  Pepsis  grossa  shows  that 
our  form  is  a  distinct  species. 

a.  NEARCTIC 
Batazoxus  interruptus  Say 

Say's  diagnosis  says  "metathorax  at  tip  bifasciate  with  yellow",  and 
the  pleura  with  two  yellow  spots.  It  came  from  Indiana.  This  is  the 
form  common  in  the  middle  states,  west  to  Kansas  and  Nebraska,  and 
in  Texas,  North  Carolina,  etc.  I  have  selected  as  neotype  a  female 
from  Chicago,  collected  by  Prof.  C.  T.  Brues,  which  agrees  with  the 
description. 

There  is  a  form  in  the  northeastern  states  which  I  call 

Batazoxus  ixterruptus  var.  cressoni  var.  now 

The  thorax  and  propodeum  are  black,  apical  margin  of  propodeum 
with  a  broadly  interrupted  yellow  line,  no  broad  yellow  band  above  it, 
no  marks  on  pleura,  the  hind  margin  of  pronotum  narrowly  yellow, 
and  the  scutellum  yellow.  The  yellow  on  each  side  of  face  in  the  typical 
form  is  here  reduced  to  a  slender  orbital  line,  the  clypeus  is  usually 
black.  The  abdomen  shows  no  pale  at  bases  of  segments,  and  the  base 
of  first  segment  is  usually  black.  The  wings  are  darker.  In  the  male 
the  yellow  is  likewise  greatly  reduced.    The  size  is  generally  smaller. 

The  holotype  is  a  female  from  Holliston,  Mass.,  4  August.  Para- 
types  from  Holliston  from  24  June  to  7  September;  Wood's  Hole, 

'Published  with  the  aid  of  a  special  gift  from  Mr.  R.  G.  Agassiz. 


168  bulletin:  museum  of  comparative  zoology 

Mass.,  (C.  T.  Brues);  Orient,  N.  Y.,  12  July;  White  Plains,  N.  Y., 
Sept.;  Wellesley,  Mass.,  13  July  (Bolster);  Trenton,  N.  J.;  Penna. 
(Melsheimer) ;  Charter  Oak,  Penna,  11  August,  (Kirk);  and  Glen- 
carlyn  and  Falls  Church  in  northern  Virginia  from  14  June  to  30  July. 

Type  M.  C.  Z.  No.  25729. 

Pompilus  ichneumonides  D.  T.  (ichneumoniformis  Patton)  and  P. 
willistoni  Patton  belong  to  Batazonus.  The  latter  I  consider  as  a  form 
of  interruptus,  and  the  former  as  B.  navus  Cress. 

Arachnophroctonus  Ferrugineus  Say 

Say  says  "Ferrugineous;  wings  violet;  pleura  and  metathorax 
black."   Inhabits  U.  S.   The  type  was  a  male. 

A  specimen  which  agrees  with  the  above  and  also  with  the  fuller 
description  I  have  selected  as  neotype;  it  is  from  Falls  Church,  Va.; 
23  July. 

There  are  three  varieties  which  can  be  separated  on  color;  there  is 
scarcely  any  difference  in  structure,  and  the  genital  plate  of  the  male 
has  a  prominent  median  carina  in  all. 

A.  ferrugineus  var.  unicolor  var.  nov. 

Some  years  ago  Viereck  labeled  a  specimen  "unicolor";  it  was  from 
the  far  west.  The  body  is  wholly  ferrugineous  except  a  small  black 
mark  at  the  base  of  the  abdomen ;  nothing  dark  on  thorax  nor  on  pro- 
podeum,  nor  even  a  trace  of  dark  bands  on  abdomen;  moreover  the 
wings  are  a  yellowish  brown,  very  much  paler  than  those  of  the  typical 
form.  The  structure  appears  to  be  the  same  except  that  the  clypeus  is 
not  so  deeply  emarginate  in  the  middle.  The  holotype  is  from  Oak 
Creek  Canon,  Arizona,  6000  ft.,  July  (F.  H.  Snow)  in  the  M.  C.  Z. 
no.  25730.  Others  are  from  Tucson,  and  southern  Arizona  (Bequaert). 

A.  ferrugineus  var.  annexus  var.  nov. 

From  parts  of  Texas  come  specimens  intermediate  in  some  ways 
between  unicolor  and  typical  ferrugineus.  There  is  no  black  on  thorax 
nor  on  propodeum,  but  the  abdomen  above  shows  at  least  traces  of  a 
dark  band  at  end  of  first  and  second  segments.  The  wings  are  just  as 
dark  as  in  typical  form,  which  separates  it  from  unicolor. 

Holotype  is  from  Fedor,  Lee  Co.,  Texas,  June  15,  1909  (Birkmann); 
paratypes  from  Fedor  in  June  and  September,  also  Dallas,  Texas 
(Boll);  Austin,  Texas,  13  October  (Brues);  and  Davis  Mts.,  Texas,  28 
June  (Englehart).   Type,  M.  C.  Z.  no.  25731. 

Typical  ferrugineus  also  occurs  in  Texas. 


hanks:  psammocharidae  169 

A.  FERRUGINEUS  var.  NIGRESCENS  var.  nOV. 

In  the  northeastern  part  of  the  country  there  is  a  much  darker  form; 
the  wings  are  nearly  black;  the  abdomen,  in  the  female,  shows  the 
dark  bands  at  tips  of  first  and  second  segments.  The  thorax  and  pro- 
podeum  are  wholly  black  above  and  below,  and  on  pleura;  the  femora 
are  at  least  partly  black,  the  hind  femora  often  wholly  so;  in  the 
female  the  tip  of  tibia  and  of  some  tarsal  joints  also  black;  the  antenna 
is  nearly  wholly  black,  basal  joint  only  remaining  pale;  in  the  male  the 
basal  segment  may  be  more  black  than  usual;  in  females  the  face 
above  the  antennae  is  more  or  less  darkened.  The  clypeus  is  less  deeply 
emarginate  than  in  typical  form;  otherwise  I  note  no  structural 
difference. 

The  holotype  is  a  female  from  Arlington,  Va.,  1 1  August;  paratypes 
from  Fort  Lee  district,  X.  J.,  August;  Woods  Hole,  Mass.,  (Brues); 
Holliston,  Mass.,  27  July  (Bks.),  and  Coldbrook,  Conn.,  14  July 
(Wheeler).   Type,  M,  C.  Z.,  no.  25732. 

Anoplius  imbellis  spec.  nov. 

Black  throughout,  wings  not  darker  at  tip.  Moderately  long  black 
hair  on  head,  thorax,  and  propodeum;  some  stiff  black  bristles  above 
tip  of  abdomen,  and  some  slender  curved  ones  above  and  below;  no 
hair  on  pleura.  ( 'lypeus  fully  two  and  one-half  times  as  broad  as  long, 
below  truncate;  antennae  slender,  third  joint  fully  five  times  as  long 
as  broad,  fourth  about  two-thirds  as  long  as  third.  Face  but  little 
narrower  above  than  below;  a  median  line  from  antennae  to  the  an- 
terior ocellus,  ocelli  subequal,  laterals  a  little  nearer  to  each  other  than 
to  eyes;  pronotum  deeply  angulate  behind;  propodeum  with  no  dis- 
tinct median  groove;  abdomen  moderately  broad  at  base,  slender  and 
pointed  at  tip;  mid  and  hind  tibiae  with  short  spines  above;  long  spur 
of  hind  tibia  nearly  three-fifths  of  basitarsus;  claws  with  short,  erect 
tooth. 

In  fore  wings  the  basal  vein  a  little  before  the  transverse,  marginal 
cell  more  than  length  from  the  wing-tip,  outer  side  straight;  second 
submarginal  cell  a  little  longer  than  broad,  receiving  the  first  recurrent 
near  tip;  third  submarginal  triangular  or  nearly  so,  outer  side  convex, 
receiving  the  second  recurrent  (lightly  curved)  near  middle. 

Length  9  7.5  to  8.5  mm. 

What  is  doubtless  the  male  is  similar,  but  little  more  slender;  the 
antennae  heavy,  third  joint  about  three  times  as  long  as  broad;  the 
third  submarginal  cell  triangular  or  even  pedicellate,  shorter  than  the 


170  bulletin:  museum  of  comparative  zoology 

second;  the  fourth  and  fifth  ventral  segments  are  covered  with  a 
brush  of  black  hair,  denser  and  longer  on  sides ;  the  genitalia  show  a 
median  plate,  narrow,  almost  pointed  at  tip,  and  with  a  median, 
scarcely  elevated  ridge. 

Length  c?  7  to  8  mm. 

Females  from  Corvallis,  Oregon,  30  August,  4  September,  and 
Hillsboro,  Oregon,  25  September. 

Males  from  Corvallis,  6,  25,  30  August,  and  Cornelius,  Oregon, 
4  August  (Scullen  coll.). 

Type  in  Oregon  Agricultural  College,  paratypes  there  and  in 
Museum  of  Comparative  Zoology  no.  25734. 

Nannopompilus  texanus  spec.  nov. 

Similar  in  structure  to  AT.  subviolaceus.  Abdomen  only  faintly 
bluish;  fore  wings  scarcely  smoky,  but  a  broad  dark  band  at  tip;  hind 
wings  hyaline,  smoky  at  tip.  The  body,  antennae,  and  basal  joints  of 
legs  sericeous,  strong  on  lower  face,  clypeus,  coxae,  and  posterior  sides 
of  propodeum.  Antennae  fully  as  short  as  in  Ar.  subviolaceus,  head, 
thorax,  and  propodeum  base  as  in  that  species.  Venation  similar,  the 
second  submarginal  cell  nearly  as  high  as  long;  the  recurrent  veins  end 
beyond  middle  of  cells.  The  legs  also  similar,  except  that  the  spines 
of  the  comb  are  longer  and  thickened,  and  some  a  little  curved,  but 
none  plainly  spatulate.  Marginal  cell  a  little  more  than  its  length  from 
tip  of  wing. 

Length  7  mm. 

One  female  from  Richmond,  Fort  Bend  Co.,  22  June,  (Bequaert) 
and  another  from  Fedor,  Lee  Co.,  (Birkmann),  both  Texas.  Type 
M.  C.  Z.  No.  25S95.  Separated  from  A:.  subviolaceus  by  the  much 
paler  wings,  sericeous  body,  stouter  spines. 

Gymnochares  texana  spec.  nov. 

Black,  pronotum  slightly  sericeous  near  hind  border,  and  in  front 
near  collar;  antennae  dark  ferruginous;  wings  slightly  darkened 
throughout,  tip  a  little  darker,  but  not  a  band.  Clypeus  more  than 
twice  as  broad  as  long,  lower  edge  truncate;  each  side  above  clypeus 
the  face  is  faintly  sericeous,  also  the  base  of  mandible;  face  with  nearly 
parallel  sides,  except  near  top;  a  very  distinct  median  groove  from  an- 
terior ocellus  to  antennae,  latter  slender,  third  joint  longer  than  ver- 
tex-width and  one  and  a  half  times  longer  than  fourth,  latter  but  little 


banks:  psammocharidae  171 

if  any  longer  than  fifth.  Hind  ocelli  a  little  nearer  the  eyes  than  to  each 
other;  vertex  with  a  few  fine,  long  hairs;  pronotum  almost  angulate 
behind,  faintly  depressed  near  hind  border,  and  with  a  median  groove 
reaching  forward;  pro  and  mesonotum  without  hair,  as  also  the  pro- 
podeum,  latter  with  a  very  distinct  median  groove  above;  on  the  sides 
of  the  posterior  slope  and  extending  over  the  side  are  distinct  but  not 
strong  transverse  ridges,  fading  out  toward  center.  Abdomen  slightly 
compressed  near  tip,  scarcely  a  trace  of  hair  above  toward  tip,  below 
a  few  near  tip. 

Legs  slender  as  in  other  species;  hind  femora  above  near  tip  with 
three  well-separated  minute  hair-pits,  rather  fainter  than  in  allied 
species;  hind  tibia  with  only  a  few  scattered  minute  spines,  three 
minute  ones  on  apical  half  of  outer  side;  inner  spur  of  hind  tibiae  not 
one-half  as  long  as  basitarsus,  latter  with  only  a  trace  of  spines  below. 

Fore  wings  with  marginal  cell  about  its  length  from  wing-tip,  about 
two  and  a  half  times  as  long  as  broad,  not  quite  as  slender  as  in  birk- 
manni;  second  submarginal  cell  fully  one  and  a  half  times  as  long  as 
broad,  receiving  the  first  recurrent  a  little  before  tip  (nearer  middle  in 
birkmanni);  third  submarginal  cell  only  a  little  longer  than  second, 
narrowed  one-third  above,  receiving  the  second  recurrent  a  little  be- 
yond middle;  basal  vein  before  transverse;  cubitus  extending  out  to 
margin  as  in  other  species.  In  hind  wing  the  anal  ends  much  before 
the  fork. 

Length  8  mm. 

One  female  from  Austin,  Texas,  May  (Melander),  Type,  M.  C.  Z. 
25704. 

Similar  in  general  to  birkmanni,  but  readily  separated  by  the  groove 
on  propodeum. 

Nannochilus  gen.  now 

Two  of  our  species  described  under  Ageniella,  externa  and  congrua, 
do  not  have  a  petiolate,  but  a  sessile  abdomen;  they  are  therefore  re- 
lated to  Priocnemis,  but  differ  from  that  genus  in  lacking  teeth  or  even 
bristles  on  the  hind  tibia.  The  claws  are  cleft,  the  inner  process  being 
very  broad  and  obliquely  truncate,  and  the  pulvillus  is  extremely 
large.  The  venation  is  similar  to  that  of  Pseudagenia,  but  the  basal 
vein  is  interstitial  with  the  transverse.  Besides  Ageniella  externa  Bks. 
which  is  the  genotype,  there  is  A.  congrua  Cress.,  and  a  new  species 
described  herewith. 


172  bulletin:  museum  of  comparative  zoology 

Nannochilus  osoria  spec.  nov. 

Body  black,  abdomen  sometimes  more  brown;  antennae  and  legs 
brown,  sometimes  the  mid  and  hind  femora  are  somewhat  rufous,  and 
the  front  tibiae  and  tarsi  are  usually  pale  yellowish,  all  spurs  pale,  but 
not  snow-white.  Clypeus  and  lower  face  with  silvery  pubescence,  little 
on  vertex,  but  more  prominent  on  coxae  and  on  each  side  of  propodeum 
behind  the  spiracles. 

Fore  wings  rather  evenly  dusky,  tip  not  darker;  veins  brown  to 
black;  hind  wings  more  hyaline. 

Clypeus  broad,  short,  and  truncate  below;  lateral  ocelli  a  little 
nearer  to  each  other  than  to  eyes;  antennae  moderately  long  and 
slender,  much  as  in  iridipennis;  pronotum  arcuate  behind,  finely 
granular,  a  median  groove  on  basal  part;  abdomen  with  short  pale  hair 
on  venter;  basal  segment  not  nearly  twice  as  long  as  broad  behind. 

Fore  wings  with  submarginal  cells  much  broader  than  the  marginal 
cell;  second  submarginal  about  as  broad  as  long,  receiving  the  first 
recurrent  near  middle;  third  submarginal  hardly  longer  below,  but 
larger,  the  posterior  side  almost  angulate  in  middle,  so  the  cell  is  no 
broader  below  than  above,  receiving  the  second  recurrent  vein  near 
middle;  basal  vein  interstitial  with  nervellus,  its  lower  part  not 
strongly  curved.  In  hind  wings  the  anal  vein  ends  before  the  fork  of 
cubitus. 

Length  of  fore  wing  5  to  6.5  mm. 

Holotype  male  from  Falls  Church,  Virginia,  12  July;  paratypes  from 
the  same  locality  from  28  June  to  7  September;  also  from  Great  Falls, 
Virginia,  20  June,  and  Chain  Bridge,  Virginia,  23  June.  Type  M.  C.  Z. 
No.  25907. 

Three  females  agree  in  general;  dull  black,  sericeous,  venation  as  in 
male,  spurs  pale,  clypeus  truncate  below;  third  antennal  joint  a  little 
longer  than  fourth;  pronotum  broadly  arcuate  behind;  inner  spur  of 
hind  tibiae  hardly  one-half  of  basitarsus;  basal  segment  of  abdomen 
broad  and  sessile. 

Length  of  fore  wing  of  female,  6  mm. 

All  from  Falls  Church,  Virginia,  13  September. 

Priocnemis  occldentis  spec.  nov. 

Closely  related  to  the  Eastern  P.  nothus;  in  color  the  wings  are  uni- 
formly darker  and  the  abdomen  is  entirely  reddish,  the  last  few  seg- 
ments not  darkened  as  in  nothus.  The  venation  the  same  as  that  of 
nothus  even  to  the  bend  in  the  lower  part  of  the  submarginal  cross- 


banks:  psammocharidae  L73 

vein;  the  propodeum  is  less  evenly  rounded  than  in  nothus,  seen  from 
the  side  there  is  almost  an  angle  at  the  turn,  the  posterior  slope  steeper 
than  in  nothus. 

The  second  plus  third  antenna]  joints  not  equal  to  vertex-width; 
ocelli  subequal,  in  a  slightly  broader  triangle  than  nothus,  the  laterals 
much  nearer  to  each  other  than  to  eyes;  pronotum  angulate  behind; 
no  median  groove  on  propodeum,  the  hue  striae  as  in  nothus;  the  spines 
and  spurs  on  legs  as  in  that  species. 

Length  6.5  to  8  mm. 

From  Corvallis,  Oregon,  20,  22,  23  July,  6,  23,  25,  30  August,  and 
4  September,  also  Forest  Grove,  1  June;  Blooming,  26  July;  and  Hills- 
boro,  24  September,  all  in  Oregon. 

Type  in  Oregon  State  College,  paratypes  there  and  in  the  M.  C.  Z. 
no.  25733. 

Priocxemis  nebulosus  Dahlb. 

Dahlbom  says  the  wings  are  violaceous,  mentions  the  emarginate 
clypeus,  and  locality  South  Carolina.  From  Florida  in  the  Graenicher 
collection  are  two  males  and  a  female  which  agree  better  with  the 
description  than  the  form  from  Virginia  northward  that,  following 
Cresson,  I  previously  identified  as  nebulosus. 

In  the  Florida  female  the  clypeus  is  more  plainly  emarginate  in  the 
middle,  the  angle  each  side  is  almost  a  tooth;  the  wings  are  more 
evenly  dark  than  the  northern  form,  which  will  be  known  as  P.  pul- 
chrina Cresson,  based  on  the  male. 

In  the  male  of  nebulosus  the  wings  are  dark  as  in  the  female  (in 
pulchrina  the  male  has  hyaline  wings  with  a  smoky  tip),  and  the  black 
median  stripe  on  the  face  is  broader,  especially  on  the  clypeus.  The 
legs  are  marked  as  in  pulchrina.  In  both  sexes  the  second  submarginal 
cell  is  not  as  long  as  in  pulchrina. 

A  female  from  Larkins,  May.  males,  South  Miami,  26  April,  and 
Lake  Apopka,  Winter  Garden,  25  April,  all  Florida. 

Priophanes 

There  are  several  among  the  described  species  of  Priocnemis  which 
have  a  distinct  petiole  connecting  the  abdomen  to  the  propodeum, 
while  true  Priocnemis  has  no  petiole.  For  the  petiolate  forms  I  pro- 
posed a  new  genus  which  is  more  related  to  Pseudagenia  and  Ageniella 
than  to  Priocnemis.  The  venation  is  much  like  Pseudagenia,  the  basal 
vein  ends  before  the  transverse,  and  in  hind  wings  the  anal  vein  ends 
before  the  cubital  fork.    The  hind  tibiae  have  three  rows  of  spines 


174  bulletin:  museum  of  comparative  zoology 

above  the  middle  one  with  more  or  less  distinct  teeth;  last  joint  of  mid 
and  hind  tarsi  bare,  the  claws  toothed;  no  distinct  "beard"  under 
head;  venter  of  female  with  a  groove  on  second  segment;  the  mesoster- 
num  not  prominent  laterally. 

Type  Priocnemis  facetus  Cress. 

Also  included  are  P.  agcnoideus,  arizonica,  arcuatus,  holonis,  n.  sp., 
placitus,  relictus. 

Priophanes  holonis  spec.  nov. 

Black;  abdomen  reddish  all  over;  wings  hyaline,  apex  broadly 
dusky. 

In  general  close  to  P.  directa  of  Texas;  the  pronotum  angulate  be- 
hind, the  spurs  black,  legs  and  antennae  entirely  black;  and  head, 
thorax,  and  abdomen  of  the  same  shape  as  P.  directa.  It  differs  in 
having  the  teeth  on  hind  tibiae  much  smaller,  hardly  noticeable,  except 
from  inner  side,  and  more  especially  in  venation.  The  marginal  cell  is 
longer,  its  outer  side  more  oblique,  and  not  quite  its  length  from  tip  of 
wing  (in  directa  more  than  its  length  from  tip  of  wing) ;  the  second  and 
third  submarginal  cells  are  proportionally  longer  than  in  directa,  and 
the  lower  outer  corner  of  third  submarginal  cell  is  scarcely  the  length 
of  third  submarginal  from  the  outer  margin  of  wing  (in  directa  more 
than  length  of  third  submarginal  cell  from  outer  margin  of  wing). 
The  first  recurrent  ends  at  or  before  middle  of  second  submarginal, 
and  second  recurrent  before  middle  of  third  submarginal  cell.  The 
basal  vein,  which  is  curved  as  in  directa,  ends  only  a  little  before  the 
transverse  vein. 

Length  7.5  mm.  to  8.5  mm. 

Holotype  from  Urbana,  Illinois,  20  July  (Bequaert);  paratypes 
Columbus,  Ohio,  August  (Bequaert),  also  6,  21  July  (Gillaspy);  and 
MacCollum,  Coweta  Co.,  Georgia,  8  June  (Bequaert).  Type  M.  C.  Z. 
25892,  paratypes  there  and  Ohio  State  University. 

Ageniella  delicata  spec.  nov. 

Dull  black,  abdomen  with  all  of  second,  most  of  first,  and  part  of 
third  segments  yellowish  rufous;  tip  of  abdomen  with  white  spot; 
mandibles  yellowish  at  tips;  antennae  brown,  basal  joint  pale  yellow 
beneath;  all  coxae  black,  rest  of  front  legs  pale  yellowish,  mid  legs 
with  tibiae  and  femora  pale,  tarsi  brown,  hind  legs  with  femora  pale, 
tibiae  and  tarsi  dark  brown;  spurs  of  hind  legs  dark  brown,  others 
paler;  wings  hyaline,  tip  not  darker,  veins  yellowish  to  brown. 


banks:  psammocharidae  175 

(  "lypeus  and  lower  face  with  silvery  pubescence;  notum  and  pleura 
somewhat  sericeous. 

Body  slender;  clypeus  broad,  somewhat  rounded  below;  lateral 
ocelli  much  nearer  each  other  than  to  eyes;  vertex  convex;  antennae 
long  and  slender,  third  joint  hardly  longer  than  fourth;  pronotum 
deeply  arcuate  behind;  propodeum  finely  granular,  median  groove 
faint;  basal  segment  of  abdomen  more  than  twice  as  long  as  broad 
behind. 

Fore  wings  rather  short,  the  submarginal  cells  not  as  broad  as  the 
marginal,  latter  nearly  its  length  before  tip  of  wing;  second  submargi- 
nal about  one-third  longer  than  broad,  receiving  the  first  recurrent 
vein  near  base;  third  submarginal  plainly  longer  on  lower  side  than 
second,  but  narrowed  nearly  one-third  above,  receiving  the  second  re- 
current (almost  angulated)  a  little  before  middle;  lower  part  of  basal 
vein  bulging  forward,  ending  a  little  before  the  nervellus. 

Mid  and  hind  legs  very  slender  and  long,  smooth,  inner  spur  of  hind 
tibiae  two-thirds  of  basitarsus. 

Length  of  fore  wing  4.2  mm. 

One  male  from  Falls  Church,  Va.,  22  August.  Type  M.  C.  Z.  No. 
25910. 

Ageniella  restricta  spec,  now 

Body  dull  black,  more  or  less  sericeous,  most  noticeable  on  the 
coxae  and  pleura;  clypeus  and  lower  face  with  dense  white  pubescence; 
abdomen  with  basal  two-thirds  of  second  segment  yellowish  rufous; 
tip  of  abdomen  with  a  white  spot;  legs  brown  to  black,  front  legs  with 
tibiae  and  tarsi  pale  yellowish,  hind  spurs  dark,  others  paler.  Fore 
wings  dusky,  tip  rather  darker,  veins  dark  brown;  hind  wings  almost 
hyaline. 

Body  slender;  clypeus  truncate  below;  antennae  slender,  third  joint 
a  little  longer  than  the  fourth,  on  the  style  of  A.  iridipcnnis;  lateral 
ocelli  plainly  nearer  each  other  than  to  eyes;  vertex  rather  strongly 
convex;  pronotum  arcuate  behind;  propodeum  finely  granular,  sloping 
toward  tip,  median  line  scarcely  visible;  abdomen  slender,  basal  seg- 
ment more  than  twice  as  long  as  broad  behind. 

In  fore  wings  the  submarginal  cells  as  broad  as  the  marginal  cell, 
latter  not  quite  its  length  from  the  tip  of  wing;  second  submarginal 
nearly  one  and  a  half  times  as  long  as  broad,  receiving  the  first  recur- 
rent before  basal  third;  third  submarginal  cell  little  longer  below  than 
second,  but  broader,  narrowed  hardly  one  third  above,  receiving  the 
second  recurrent  (nearly  evenly  curved)  before  middle:  lower  part  of 


176  bulletin:  museum  of  comparative  zoology 

basal  vein  only  slightly  bulging,  and  ending  a  little  before  the  nervel- 
lus;  mid  and  hind  legs  long  and  smooth,  inner  spur  of  hind  tibia  more 
than  one-half  but  hardly  two-thirds  of  basitarsus. 

Length  of  fore  wing  5  mm. 

One  male  from  Falls  Church,  Virginia,  22  August,  on  leaves  of 
tulip-tree  with  honey-dew.  Type  M.  C.  Z.  No.  25909.  Closely  re- 
lated to  A.  delicata,  but  differing  in  less  reddish  on  abdomen,  basal 
joint  of  antennae  black,  shorter  spurs,  darker  wings  and  venation,  and 
broader  submarginal  cells. 

Ageniella  neglecta  spec.  nov. 

Body,  antennae,  legs,  wholly  black,  hind  spurs  black,  others  paler; 
fore  wings  nearly  hyaline,  slightly  darker  near  tip,  veins  brown. 
Clypeus  and  lower  face  with  white  pubescence,  thorax  and  coxae 
somewhat  sericeous,  not  on  abdomen. 

Clypeus  truncate  below;  antennae  short  and  rather  thick,  the  third 
joint  little  more  than  twice  as  long  as  broad  at  tip ;  pronotum  broadly 
arcuate  behind;  propodeum  (from  side)  slightly  rounded,  no  median 
line;  basal  segment  of  abdomen  not  quite  twice  as  long  as  broad  be- 
hind. Legs  short,  smooth,  inner  spur  of  hind  tibiae  little  more  than 
one-half  of  basitarsus. 

In  fore  wings  the  marginal  cell  is  fully  its  length  before  tip  of  wings, 
the  third  submarginal  cell  nearly  three  times  its  length  from  margin, 
and  almost  as  broad  as  the  marginal  cell;  second  submarginal  cell  a 
little  longer  than  broad,  receiving  the  first  recurrent  at  basal  third; 
third  submarginal  a  little  longer  than  the  second,  only  a  little  nar- 
rowed above,  receiving  the  slightly  curved  second  recurrent  near  tip; 
lower  part  of  basal  vein  only  a  little  convex  and  ends  on  the  nervellus ; 
in  hind  wings  the  anal  ends  much  before  cubital  fork. 

Length  of  fore  wing  3.5  mm. 

One  male  from  Boulder,  Colorado,  26  August,  1908  (S.  A.  Rohwer). 
Type  M.  C.  Z.  No.  25908. 

Readily  known  by  small  size,  deep  black  color,  short  legs,  and  the 
large  apical  field  beyond  venation. 

Ageniella  accepta  var.  conflicta  var.  nov. 

Differs  from  the  typical  form  in  having  a  black  spot  on  side  of  scutel- 
lum  and  on  side  of  metanotum  as  well  as  at  base  of  fore  wing,  usually 
all  run  together  in  a  black  stripe;  area  between  ocelli  black;  body  more 


banks:  psammochajridae  177 

slender  than  typical  form;  the  pale  area  of  fore  wing  between  the  first 
brown  band  and  the  stigma]  band  is  more  suffused  with  pale  brown,  so 
that  the  wing  is  not  so  plainly  three-banded;  the  basal  part  of  wing 
is  more  yellowish,  and  the  brown  at  apex  is  a  paler  brown  than  in 
typical  acccpta. 

Holotype,  a  female  from  Falls  Church,  Va.,  5  July;  paratypes  also 
from  Falls  Church  from  19  June  to  13  Sept.;  from  Glencarlyn,  Va., 
26  July,  and  from  Riverhead,  L.  I.,  X.  Y.,  1  August  (W.  T.  Davis). 
Type  M.  C.  Z.  no.  25735. 


Synoptic  table  of  males  of  eastern  species 
of  Ageniella  at  present  known  to  me 

1.  Tip  of  abdomen  with  a  distinct  white  spot;  third  cubital  cell  longer  below 

than  above 2 

Tip  of  abdomen  wholly  dark 11 

2.  Some  reddish  or  yellowish  on  dorsum  of  abdomen;  basal  segment  of  abdo- 

men more  than  twice  as  long  as  broad 3 

No  reddish  nor  yellowish  on  dorsum  of  abdomen 7 

3.  All  spurs  snow-white ;  femora  rufous birkmanni 

At  least  hind  spurs  dark 4 

4.  Clypeus  with  a  small  pale  spot  on  each  side,  and  a  small  orbital  spot  .festina- 
Clypeus  wholly  dark 5 

5.  A  small  yellow  orbital  line  or  spot;  marginal  cell  much  less  than  its  length 

from  tip  of  wing;  inner  spur  of  hind  tibiae  scarcely  more  than  one-half  of 

basitarsus;  basal  joint  of  antennae  dark fratemella 

Marginal  cell  nearly  its  length  from  tip  of  wing;  no  pale  orbital  line  or  spot; 
inner  spur  of  hind  tibiae  more  than  one-half  of  basitarsus 6 

6.  Basal  joint  of  antennae  pale  yellowish  below,  femora  yellowish  or  rufous, 

paler  than  rest  of  legs;  red  of  abdomen  on  all  of  second  and  parts  of  first 

and  third  segments delicata 

Basal  joint  of  antennae  black  below;  femora  more  brown,  darker  than  rest 
of  legs;  red  on  abdomen  only  on  part  of  second  segment restricta 

7.  Two  pale  spots  or  a  white  band  on  hind  border  of  pronotum ;  clypeus  mostly 

whitish;  spurs  snow-white calcarata 

No  pale  mark  on  pronotum 8 

8.  A  small  yellowish  spot  or  short  stripe  on  orbital  line  a  little  above  clypeus; 
long  spur  of  hind  tibiae  about  one-half  of  basitarsus;  vertex  usually  shin- 
ing; fore  wings  more  than  five  millimeters  long agilis 

No  such  spot  or  stripe  on  orbital  line;  fore  wings  not  over  five  millimeters 
long 9 


178  bulletin:  museum  of  comparative  zoology 

9.  Femora  rufous  or  yellowish,  sometimes  tibiae  also;  inner  spur  of  hind 

tibiae  hardly  more  than  half  of  basitarsus texana 

Femora  and  tibiae  black;  inner  spur  of  hind  tibiae  more  than  one-half  of 
basitarsus 10 

10.  Abdomen  very  slender,  second  and  third  segments  with  pendent  side-mem- 

brane, basal  segment  plainly  more  than  twice  as  long  as  broad  behind. 

pctiolata 

Abdomen  broader,  no  side-membrane  to  second  and  third  segments;  basal 

segment  twice  as  long  as  broad  behind,  tip  of  fore  wing  black norata 

11.  Abdomen  with  some  reddish  or  yellowish  on  basal  half  above 12 

Abdomen  wholly  black  above 14 

12.  Basal  segment  of  abdomen  not  twice  as  long  as  broad  behind;  third  sub- 

marginal  cell  with  angulate  hind  border perfecia 

Basal  segment  of  abdomen  more  than  twice  as  long  as  broad;  small  species. 

13 

13.  Clypeus  black;  second  segment  of  abdomen  dark  in  middle,  yellowish  on 

sides;  third  submarginal  cell  higher  than  long;  hind  spurs  dark;  basal 
segment  of  antennae  black;  third  submarginal  angulate  behind. 

minuscula 

Clypeus  mostly  pale;  spurs  white;  basal  joint  of  antennae  pale  above  and 

below;  hind  border  of  third  submarginal  cell  not  at  all  angulate  behind. 

apicipennis 

14.  Clypeus  with  some  pale,  at  least  on  lower  edge;  palpi  pale 15 

Clypeus  wholly  dark 17 

15.  Third  submarginal  cell  with  outer  border  angled  in  middle,  so  cell  no 

broader  below  than  above;  front  coxae  almost  white clypeata 

Third  submarginal  cell  plainly  longer  on  lower  side  than  on  upper 16 

16.  Hind  border  of  pronotum  slightly  pale;  basal  joint  of  antennae  white  below. 

hestia 
Hind  border  of  pronotum  shows  no  paler  band;  basal  segment  of  abdomen 
not  twice  as  long  as  broad  behind;  antennae  thicker  than  usual. 

crassicornis 

17.  Third  submarginal   cell  with  hind  border  angled  in  middle,  so  cell  no 

broader  below  than  above 18 

Third  submarginal  cell  with  outer  side  sloping,  and  lower  border  longer 
than  upper 19 

18.  Hind  spurs  dark;  second  recurrent  arises  near  the  middle  of  outer  part  of 

cubitus;  small  species eximia 

All  spurs  nearly  white;  second  recurrent  arises  plainly  beyond  the  middle  of 
outer  part  of  cubitus ;  larger  species osoria 

19.  All  spurs  snow-white;  basal  segment  of  abdomen  fully  twice  as  long  as 

broad  behind virginica 

Basal  segment  not  quite  twice  as  long  as  broad  behind;  all  spurs  not  white 

20 


banks:  psammocharidae  17!) 

20.  Third  suhmarginal  cell  much  longer  than  high;  a  distinct  median  groove 

on  propodeum species? 

Third  submarginal  cell  scarcely  as  long  as  high,  often  plainly  higher  than 
long  below;  no  distinct  groove  on  propodeum 21 

21.  Hind  femora  reddish  (or  yellowish)  on  outer  half,  base  black  or  brown. 

aludra 
If  hind  femora  pale  it  is  also  on  basal  part 22 

22.  Second  recurrent  arises  little  if  any  beyond  middle  of  outer  part  of  cubitus. 

23 
Second  recurrent  arises  plainly  beyond  middle  of  outer  part  of  cubitus .  .  24 

23.  Front  femora,  tibiae,  and  tips  of  coxae  pale;  third  submarginal  cell  always 

longer  below  than  above;  hind  legs  often  partly  pale tenella 

Front  femora,  tibiae,  and  coxae  not  pale;  third  submarginal  cell  often  no 
longer  below  than  above;  legs  black,  a  small  species eximia 

24.  Front  tarsi  yellowish;  basal  segment  of  abdomen  about  one  and  one-half 

times  as  long  as  broad  behind iridipennis 

Front  tarsi  not  yellowish;  basal  segment  of  abdomen  not  one  and  one-half 
times  as  long  as  broad  behind atrata 

Nemagenia  subgenus  nov.  of  Ageniella 

Pronotum  above  as  long  as  mesonotum ;  marginal  cell  its  length  from 
tip;  basal  vein  plainly  before  nervellus;  inner  spur  of  hind  tibiae  not 
one-half  of  basitarsus;  propodeum  very  long,  nearly  flat,  from  side 
it  is  only  slightly  curved;  otherwise  like  Ageniella. 

Type  Pompilus  (Agenda)  longulus  Cresson. 

Only  the  male  is  known;  described  from  North  Dakota,  I  have 
specimens  from  Fedor,  Texas  (Birkmann). 

Dipogon  texanus  spec.  nov. 

Head  and  abdomen  black,  clypeus  and  thorax  rufous,  legs  dull 
black  to  brown,  tarsi  partly  paler,  coxae  rufous,  antennae  yellowish, 
a  narrow-black  band  at  tip  of  each  joint,  except  basal  and  apical;  fore 
wings  hyaline,  a  narrow  black  band  across  over  the  basal  vein,  and  a 
very  broad  blackish  band  across  over  submarginals  and  third  discoidal 
cells,  not  occupying  the  tip  of  marginal  cell. 

Face  and  vertex  with  short  appressed  white  hair ;  abdomen  also  with 
pale  appressed  hair,  longer  and  erect  hair  near  tip;  venter  with  bands 
of  pale  hair.  Hair  basket  under  head  of  white  bristles.  Structure  in 
general  similar  to  D.  brevis;  very  little  hair  on  thorax  or  propodeum; 
venation  similar,  but  marginal  cell  hardly  as  broad,  but  angulate  on 
hind  border;  second  submarginal  about  twice  as  long  as  the  third;  first 


180  bulletin:  museum  of  comparative  zoology 

recurrent  ending  much  before  middle  of  second  submarginal,  second 
recurrent  ending  near  base  of  third  submarginal  cell;  medius  reaches 
margin;  legs  slender  as  usual;  long  spur  of  hind  tibia  about  two-fifths 
of  basitarsus. 

Length  6.5  mm. 

From  Brownsville,  Texas,  11  to  16  June  (Darlington).  Type 
M.  C.  Z.  No.  25896. 

Dipogon  sericea  spec.  nov. 

Body  black;  antennae  with  first  and  second  joints  black,  beyond 
yellowish,  some  joints  narrowly  dark  at  tips;  legs  black,  tarsi  and  the 
front  tibiae  rufous.  Fore  wings  brown,  beyond  the  marginal  cell 
snow-white,  the  marginal  cell  and  below  and  beyond  it  darker  than 
elsewhere,  stigma  black;  hind  wings  slightly  infuseate. 

Thorax  and  abdomen  densely  clothed  with  appressed  gray  pu- 
bescence, in  places  somewhat  yellowish,  thorax  with  rather  long,  erect 
white  hairs,  and  shorter  ones  on  the  abdomen,  near  tip  darker.  Face 
above  antennae  with  bright  yellowish  pubescence,  across  face  half  way 
up  to  vertex  is  a  band  of  erect  black  bristles,  and  similar  bristles  on 
vertex;  the  lower  edge  of  clypeus  is  yellow,  rest  black,  clothed  with 
white  hairs,  white  hairs  back  of  eyes;  hair  basket  of  fine,  pale  bristles. 

Legs  with  the  femora,  mid  and  hind  tibiae  covered  with  sericeous 
pile,  the  femora  with  fine,  long  hairs  below.  Hind  tibiae  above  with  a 
groove  and  a  row  of  fine  short  hairs,  long  spur  about  one-third  of  the 
basitarsus.  Antennae  rather  slender,  third  joint  not  much  longer  than 
fourth,  together  they  equal  vertex-width. 

Thorax  shorter  than  in  Eastern  species,  but  fully  as  broad ;  pronotum 
broadly  arcuate  behind;  propodeum  appears  to  have  a  broad  median 
furrow,  but  all  covered  with  the  appressed  hair. 

In  fore  wings  the  venation  is  much  like  the  Eastern  D.  sayi  except 
that  the  marginal  cell  is  much  shorter,  extending  only  a  trifle  beyond 
the  third  submarginal  cell;  the  medius  reaches  to  the  margin,  but  in 
the  white  area  it  is  very  fine.  In  hind  wings  venation  as  in  other 
species. 

Length  of  fore  wing  6  mm.,  body  6.5  mm. 

One  from  Bull  Prairie,  Lake  Co.,  Oregon,  22  July,  Camas  Prairie 
Summit,  7,500  ft.  (Frewing  coll.)   Type  at  Oregon  State  College. 

The  species  of  Dipogon  so  far  described  from  the  United  States 
can  be  separated  by  the  following  key : 

1.  The  medius  of  fore  wing  plainly  does  not  reach  the  margin;  marginal  cell  not 
strongly  angled  at  end  of  second  submarginal  cell,  the  outer  side  curved. 

Subgenus  Adipogon — 2 


banks:  psammocharidae  181 

The  medius  reaches  to  the  outer  margin;   the  marginal   cell    is   strongly 
angled  at  the  end  of  the  second  submarginal  cell,  and  broadest  at  this  point. 

Subgenus  Dipoyon — 3 

2.  Forewings  wholly  dark,  nearly  black papaya 

Forewings  clear  with  a  narrow  dark  band  over  the  basal  and  transverse 

veins,  and  a  large  dark  spot  over  marginal  cell  and  the  two  or  more  cells 
behind  it pulchripennis 

3.  Thorax  yellowish  to  rufous 4 

Thorax  black 5 

4.  Head  and  abdomen  black;  antennae  rufous,  tips  of  joints  narrowly  black; 

legs  partly  black texan  us 

Head  rufous,  abdomen  black;  antennae  rufous,  no  dark  bands;  legs  wholly 
yellowish yraenicheri 

5.  Face,  mesonotum,  and  abdomen  with  appressed  grayish  to  yellowish  pu- 

bescence; hair  basket  pale;  front  legs  partly  pale 6 

Face,  thorax,  and  abdomen  without  such  pubescence;  fore  wings  clear  with 
two  bands 7 

6.  Across  face  and  on  vertex  are  erect  black  bristles;  fore  wings  brown,  tip 

snow-white;  propodeum  and  pleura  also  sericeous sericea 

No  such  bristles  on  face  and  vertex;  extreme  wing-tip  faintly  dark,  no  snow- 
white;  propodeum  and  pleura  without  appressed  hair brevis 

7.  Front  legs  largely  yellowish,  also  mandibles,  and  lower  edge  of  clypeus. 

caliptera 
Front  legs,  clypeus,  and  mandibles  black sayi 

Adipogon,  subgenus  nov.  has  the  basket  of  curved  hairs  on  under 
side  of  head  as  in  typical  Dipogon,  but  the  median  vein  of  fore  wing 
does  not  extend  to  the  outer  margin,  and  the  marginal  cell  is  not 
angled  below. 

Type  is  Pompilus  pulchripennis  Cresson. 

Pepsis  pattoni  spec.  nov. 

Our  large  black  Arizona  Pepsis  has  been  identified  by  Fox  and  my- 
self as  P.  obliquerugosa  Lucas,  a  form  described  from  Cuba.  A  few 
years  ago  Salman  considered  both  to  be  P.  grossa  Fabr.  of  northern 
South  America.  With  a  series  of  both  sexes  of  the  three  forms  I  con- 
sider each  a  separate  species. 

As  with  the  others,  the  wings  are  black  with  a  narrow  pale  band  at 
tip,  not  as  broad,  and  in  the  female  not  as  white  as  in  obliquerugosa. 
On  the  propodeum  the  side  tubercles  are  very  prominent,  sometimes 
almost  pointed,  in  obliquerugosa  much  lower  and  less  noticeable.    At 


182  bulletin:  museum  of  comparative  zoology 

the  middle  of  the  turn  there  is  in  both  species  a  high  ridge;  in  obliqueru- 
gosa  its  top  is  straight  or  a  little  convex,  in  pattoni  the  top  is  plainly 
emarginate  in  the  middle.  The  oblique  ridges  on  the  propodeum  so 
characteristic  of  obliqucrugosa,  are  fewer,  less  oblique,  and  sometimes 
none  oblique.  In  pattoni  the  third  joint  of  antennae  in  female  is  a 
little  shorter  than  in  obliqucrugosa  in  specimens  of  same  wing-length. 
In  the  male  the  parameres  of  the  genitalia  are  slender  and  tapering 
toward  tip;  in  grossa  (see  Lucas  figure  24)  they  are  heavier,  and  not  at 
all  tapering;  in  obliqucrugosa  the  parameres  are  not  as  broad  as  in 
grossa,  but  do  not  taper  as  much  as  in  pattoni;  in  the  latter  the  tip  of 
the  subgenital  plate  is  broadly  convex,  in  the  Cuban  form  it  is  plainly 
emarginate  in  middle.  The  females  of  all  three  forms  have  long,  stiff, 
curved  bristles  under  the  front  femora,  and  with  our  ncphclc  and  a  few 
South  American  species  they  form  a  natural  group.  Lucas  placed  the 
male  of  ncphclc  asformosa  and  says  that  the  genitalia  are  the  same  as  in 
grossa,  and  gives  no  separate  figure.  The  males  of  this  section  have 
the  subgenital  plate  slender  and  furnished  with  a  broad  dense  crest  of 
long  black  hairs. 

P.  obliqucrugosa  is  somewhat  the  largest  of  the  three  species,  fore 
wing  sometimes  45  mm.  P.  pattoni  sometimes  about  42  mm.,  fre- 
quently 40  mm.  P.  grossa  rarely  40  mm.,  often  near  35  mm. 

Holotype  of  P.  pattoni  9  is  from  Palmerlee,  Arizona,  July  (Bieder- 
mann  coll.),  M.  C.  Z.  No.  25805;  allotype  from  southern  Arizona; 
paratypes  are  from  Palmerlee,  Tucson,  Ola,  Ft.  Grant,  Pinalerro 
Mts.,  Santa  Catalina  Mts.,  all  Arizona,  from  late  June  to  September, 
taken  by  Snow,  Bequaert,  Morse,  and  Wheeler.  One  from  mountains 
near  Pomona,  California,  by  H.  C.  Fall. 


b.  ANTILLEAN 

Batazonus  gundlachi  Cress. 

Described  from  female  only. 

Male.  It  is  very  similar  to  female,  and  marked  the  same;  thorax 
and  abdomen  brownish  red,  the  clypeus  is  reddish  in  middle,  above 
antennae  are  two  short  curved  dark  streaks,  there  is  some  yellow  each 
side  of  scutellum  (as  well  as  the  post-scutellum) ;  a  large  yellow  spot 
just  above  base  of  mid  coxae;  propodeum  broadly  yellow  across  apical 
half;  abdomen  black  at  extreme  base,  and  faintly  dark  across  tip  of 
first  segment,  a  broad  yellow  band  across  base  of  third  segment  above 
and  below;  legs  reddish,  except  the  paler  tarsi  which  are  dark  at  tips 


banks:  psa mmoch arid ae  183 

of  joints;  inner  spur  of  hind  tibia  about  three-fifths  of  basitarsus.  The 
third  submarginal  cell  is  much  longer  than  high,  as  in  the  female. 

Length  of  fore  wing  13  mm. 

Allotype  from  San  Bias,  Trinidad  Mts.,  Cuba,  24  April  (G.  E.  Folk). 

Batazonus  hookeri  Rohwer 

Described  from  female  only. 

The  male  is  largely  yellowish;  there  is  a  triangular  black  spot  below 
antennae,  two  broad  black  stripes  above,  narrowly  separated;  two 
yellow  lines  on  mesonotum,  the  mesopleura  has  two  slightly  separated 
elongate,  yellow  spots;  the  propodeum  above  is  yellow,  black  at  ex- 
treme base,  usually  extended  back  in  middle  to  divide  or  partly  divide 
the  yellow.  The  dark  bands  on  abdomen  above  are  brown  to  black, 
occupying  hardly  one-half  of  the  segment,  basal  segment  dark  only  at 
tip;  coxae  black,  with  a  yellow  mark;  femora  partly  dark,  tip  pale;  tips 
of  tarsal  joints  black;  inner  spur  of  hind  tibiae  about  four-fifths  of  the 
basitarsus. 

Length  of  fore  wing  5  to  7  mm. 

Allotype  from  Hatillo,  Puerto  Rico,  January ;  others  from  Mayaguez 
in  November  and  Cartagena  in  May,  both  Puerto  Rico,  all  through 
Mr.  Ramos. 

PSAMMOCHARES  PARSONSI  Spec.  nOV. 

Body,  legs,  and  antennae  wholly  deep  black;  wings  mostly  black,  but 
paler  in  third  discoidal  cell  and  toward  base  both  in  front  and  behind; 
hind  wings  strongly  fumose. 

Clypeus  over  three  times  as  broad  as  long,  nearly  truncate  below, 
a  few  hairs  on  lower  part;  ocelli  in  a  low  triangle,  the  laterals  a  little 
nearer  to  the  eyes  than  to  each  other;  a  median  line  above  base  of 
antennae,  latter  slender,  third  joint  equal  vertex-width,  last  joint  only 
about  one-half  of  third  joint;  vertex  with  a  few  long,  erect  hairs, 
shorter  black  hair  on  front. 

Pronotum  broadly  arcuate  behind;  mesonotum  with  a  few  long  hairs 
each  side,  pleura  with  only  scattered  short  hairs;  propodeum  short,  no 
distinct  median  groove,  with  rather  short  hair  above. 

Abdomen  hairy  toward  tip  below,  above  with  stiff  bristles  just  be- 
fore tip,  otherwise  a  few  hairs  on  venter  near  hind  border  of  segments. 

Femora  smooth,  without  hairs;  front  tarsi  with  a  distinct,  though 
short  comb,  a  spine  at  middle  of  the  second  tarsal  joint;  hind  tibiae 
quite  heavily  spined;  two  irregular  rows  above,  some  as  long  as  the 


184  bulletin:  museum  of  comparative  zoology 

width  of  the  joint;  inner  spur  of  hind  tibia  more  than  one-half  of 
basitarsus. 

In  fore  wings  the  marginal  cell  is  about  its  length  before  tip ;  second 
submarginal  cell  longer  below  than  high,  one-third  narrowed  above, 
receiving  the  first  recurrent  vein  near  tip;  third  submarginal  cell  only 
a  little  longer  than  second  and  likewise  narrowed  one-third  above, 
receiving  the  second  recurrent  vein  (evenly  curved)  beyond  the 
middle.    In  hind  wings  the  anal  ends  beyond  the  fork. 

Length  of  fore  wing  10  mm. 

Females  from  Buenos  Aires,  Trinidad  Mts.,  Cuba,  17  to  23  June 
(C.  T.  Parsons). 

In  appearance  like  P.  pcrpilosus  but  easily  separated  by  absence  of 
hairs  on  femora  and  very  much  less  hair  on  body.  Type  M.  C.  Z. 
No.  25742. 

NOTIOCHARES  ANTILLANA  spec.  nov. 

Male.  Very  much  like  AT.  cvbensis,  wholly  black,  with  blue  and 
violet  reflections,  and  agreeing  closely  in  structure  to  that  species. 
The  only  difference  of  importance  is  that  the  male  genital  plate,  which 
is  N .  cubensis  ends  in  a  sharp-pointed  tooth  each  side,  in  N.  antiUana 
ends  in  short,  rounded  lobe,  alike  in  all  three  specimens. 

Length  of  fore  wing  8  to  11  mm. 

From  Barbados,  August  to  December  (Spencers).  Type  M.  C.  Z. 
No.  25743. 

Episyron  cressoni  Dewitz 

The  male  is  black;  a  white  spot  on  each  side  of  clypeus;  a  broad 
orbital  streak  yellowish  white,  reaching  nearly  to  vertex;  a  narrow 
yellowish  band  across  tip  of  pronotum,  a  white  spot  each  side  on  basal 
part  of  third  abdominal  segment,  the  last  dorsal  segment  somewhat 
rufous,  but  white  at  tip;  the  mid  femora  rufous  except  base  and  tip, 
the  hind  femora  rufous  except  basal  third,  and  the  hind  tibiae  rufous 
except  extreme  tip,  spurs  are  pale;  inner  one  of  hind  tibia  nearly  equal 
to  basitarsus. 

Length  of  fore  wing  6  mm. 

Allotype  from  Mona  Island,  April  (Ramos). 

Priocnemis  ursula  spec.  nov. 

Body  black,  much  of  it  covered  with  a  fine  sericeous  pile  or  bloom; 
clypeus,  mandibles,  basal  joint  of  antennae,  and  a  few   joints  beyond 


banks:  psammocharidae  IS.") 

yellowish,  rest  of  antennae  brown;  legs  black;  wings  hyaline,  venation 
black. 

Clypeus  above  about  as  broad  as  vertex,  nearly  evenly  rounded 
below,  fully  two  and  one-half  times  as  broad  as  long,  with  a  few  fine 
hairs;  front  with  fine,  pale  hairs,  and  vertex  with  some  longer  ones; 
ocelli  in  nearly  equilateral  triangle,  laterals  a  little  (but  not  much) 
nearer  to  each  other  than  to  the  eyes;  a  short  median  groove  above 
base  of  antennae,  latter  slender,  second  plus  third  joints  not  as  long 
as  vertex-width;  pronotum  scarcely  constricted  behind,  with  a  broadly 
arcuate  hind  margin ;  mesonotum  not  humped  in  middle,  scarcely  hairy, 
scutellum  and  metanotal  lobe  with  erect  black  hair;  propodeum  with 
rather  long,  fine,  pale  hairs  above. 

Abdomen  hairy  only  toward  tip  and  somewhat  beneath;  legs 
moderately  slender,  hind  tibia  above  with  about  ten  teeth  with  a 
short  spine,  and  a  row  of  spines  on  outer  part;  inner  spur  of  hind  tibiae 
two-fifths  of  basitarsus. 

The  fore  wings  have  the  marginal  cell  rather  longer  than  space  to  the 
tip  of  wing,  three  times  as  long  as  broad,  outer  side  nearly  straight; 
second  submarginal  cell  oblique,  below  nearly  twice  as  long  as  high, 
receiving  the  first  recurrent  vein  beyond  the  middle;  the  third  sub- 
marginal  cell  plainly  longer  than  second,  but  not  as  much  wider  as  in 
many  species,  outer  margin  bent  near  median  vein,  receiving  the 
second  recurrent  vein  (almost  straight)  much  before  middle;  in  hind 
wings  the  anal  vein  ends  much  before  the  fork. 

Length  of  fore  wing  6.5  mm. 

Female  from  Villa  Altagracia,  San  Domingo,  July  (Darlington) 
Type  M.  C.  Z.  No.  25741. 

In  general  very  similar  to  P.  salti  Bks.  which  differs  in  the  yellowish 
tip  to  the  abdomen. 

Priocnemis  arioles  spec.  nov. 

Head  and  thorax  black,  abdomen  reddish  except  black  tip,  the  fifth 
segment  above  bluish,  thoracic  notum  bluish,  legs  black,  femora  and 
tibiae  somewhat  bluish;  antennae  black;  fore  wings  nearly  evenly 
infumate,  slightly  darker  toward  costal  tip  than  behind,  hind  wings 
slightly  smoky. 

Clypeus  no  wider  than  face  below,  with  a  few  pale  hairs,  about  two 
and  one-half  times  as  broad  as  long,  truncate  below;  vertex  as  broad 
as  face  below,  scarcely  convex,  with  a  few  erect  hairs  each  side,  none 
on  front;  ocelli  in  a  rather  low  triangle,  the  laterals  much  nearer  each 


186  bulletin:  museum  of  comparative  zoology 

other  than  to  eyes ;  a  groove  from  anterior  ocellus  to  base  of  antennae. 

Pronotum  plainly  contracted  behind,  the  margin  slightly  arcuate; 
mesonotum  almost  humped  in  middle,  with  one  or  two  erect  bristles 
each  side  and  a  pair  on  scutellum,  pleura  bare;  propodeum  with  only 
faint,  erect,  pale  hair  above,  a  whitish  pile  across  base,  and  an  elongate 
spot  each  side  toward  hind  margin,  surface  minutely  granulate,  no 
median  groove.  Abdomen  hairy  on  venter  and  apical  third  above. 
Legs  not  especially  slender,  mid  tibiae  with  two  rows  of  short  spines 
above,  hind  tibiae  with  a  row  of  seven  or  eight  oblique  teeth,  a  spine 
beyond  each,  and  an  outer  row  of  short  spines ;  inner  spur  of  hind  tibia 
hardly  two-fifths  of  basitarsus. 

Marginal  cell  of  fore  wings  about  its  length  from  wing-tip,  two  and 
one-half  times  as  long  as  broad,  outer  side  convex;  second  submarginal 
cell  a  little  longer  than  high,  but  little  narrowed  above,  receiving  the 
first  recurrent  vein  at  middle ;  third  submarginal  cell  much  longer  than 
second  and  much  wider  behind,  one-third  narrowed  above,  receiving 
the  second  recurrent  vein  (slightly  curved)  a  little  before  middle;  in 
hind  wings  the  anal  vein  ends  much  before  the  fork. 

Length  of  fore  wing  6  mm. 

Female  from  Constanza,  Valle  Nuevo,  San  Domingo,  21  August, 
3-  to  4000  ft.  (Darlington).  Type  M.  C.  Z.  No.  25740. 

The  Antillean  species  with  a  more  or  less  reddish  abdomen  can  be 
tabulated  as  follows : — 

1.  Fore  wings  hyaline  with  two  broad  dark  bands pulchellus 

Fore  wings  without  bands 2 

2.  Thorax  wholly  reddish christophei 

Thorax  black 3 

3.  Third  submarginal  cell  no  longer  than  second,  higher  than  long parous 

Third  submarginal  cell  much  longer  than  second,  and  much  longer  than 

high 4 

4.  Wings  hyaline;  hind  legs  very  long,  body  about  4.5  mm dowi 

Wings  plainly  infuscate,  hind  legs  not  so  long,  body  7  mm ariolcs 

Priocnemella  domingensis  spec.  nov. 

Head,  propodeum,  and  pleura  black,  thorax  and  abdomen  above 
blue.  Fore  wings  black,  violaceous,  hind  wings  infuscated,  also  with 
violet  reflections;  antennae  and  legs  black,  the  femora  and  tibiae 
above,  and  the  front  coxae  iridescent  bluish,  also  basal  joint  of  anten- 
nae above. 

Clypeus  large,  extending  laterally  under  the  eyes,  with  long,  black 


banks:  psammocharidae  187 

hairs,  lower  margin  narrowly  smooth  and  coming  to  a  blunt  point  in 
middle;  front  and  vertex  with  long  black  hair,  one  each  side  on  vertex 
very  long;  third  antennal  joint  equal  to  vertex-width.  Ocelli  close  to- 
gether, the  laterals  more  than  twice  as  close  to  each  other  as  to  the 
eyes;  groove  from  anterior  ocellus  to  the  antennae.  Scattered  erect 
black  hairs  on  notum  and  scutellum,  short  hair  on  pleura;  pronotum 
almost  angled  behind.  Propodeum  minutely,  transversely  striate, 
more  distinctly  so  than  in  P.  violaceipes,  above  and  on  sides  with 
sparse,  erect  black  hair. 

Abdomen  hairy  near  tip  and  below,  scarcely  so  near  base.  Legs 
slender,  spined  as  in  P.  violaceipes,  the  mid  and  hind  tibiae  with  two 
rows  of  erect,  short  black  spines,  no  teeth,  and  evenly  short  spines  on 
the  tarsal  joints;  claws  with  an  erect,  small  tooth;  long  spur  of  hind 
tibia  about  two-fifths  of  the  basitarsus. 

Venation  of  wings  as  in  P.  violaceipes;  the  marginal  cell  long  and 
pointed;  second  submarginal  oblique,  about  one  and  one-half  times  as 
long  as  high,  receiving  the  first  recurrent  vein  near  middle;  the  third 
submarginal  cell  much  longer,  about  one-third  narrowed  above,  re- 
ceiving the  second  recurrent  also  near  middle,  this  recurrent  is  more 
sinuously  curved  than  that  in  P.  violaceipes. 

Length  of  fore  wing  10.5  to  11  mm. 

Two  females  from  near  and  southeast  of  Constanza,  Valle  Nuevo, 
San  Domingo,  August,  3  to  7000  ft.  (Darlington).  Type  M.  C.  Z.  No. 
25739. 

Except  for  the  striking  difference  in  color  of  thorax  and  abdomen, 
and  slight  differences  in  structure,  it  is  practically  the  same  as  P.  vio- 
laceipes of  Cuba. 

Priochilus 

A  few  Neotropical  species  have  much  the  appearance  of  Priocnemis; 
however,  the  last  joint  of  hind  tarsi  has  spines  beneath,  but  not  later- 
ally; the  hind  tibia  has  no  real  teeth  like  Priocnemis,  but  spines  in 
rows, •some  as  long  as  the  diameter  of  the  joint ;  the  claws  are  cleft,  the 
inner  part  broader  than  the  outer  part,  and  somewhat  obliquely  trun- 
cate; the  palpi  are  slender  as  in  Priocnemis. 

Pompilus  nobil is  Fabr.  is  the  genotype;  Salius  opacifrons  Fox  from 
Jamaica  goes  in  the  genus,  although  smaller  than  most  others.  In 
South  America  there  are  regius  Fabr.,  diversus  Smith,  scrupulus  Fox, 
sericeifrons  Fox  and  others.  Mr.  Williams  in  his  paper  "Studies  in 
Tropical  Wasps",  192S,  p.  141,  calls  attention  to  nobilis  and  regius  as 
possibly  forming  a  new  genus. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT    HARVARD    COLLEGE 

Vol.  XCIV,  No.  5 


SCIENTIFIC  RESULTS  OF  A  FOURTH  EXPEDITION 
TO  FORESTED  AREAS  IN  EAST  AND  CENTRAL  AFRICA 

VI 
ITINERARY  AND  COMMENTS 


By  Arthur  Loveridge 


With  Four  Plates 


CAMBRIDGE,  MASS.,  U.S.A. 

PRINTED    FOR    THE    MUSEUM 

July,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2, 
3  and  4  have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT    HARVARD    COLLEGE 

Vol.  XCIV,  No.  5 


SCIENTIFIC  RESULTS  OF  A  FOURTH  EXPEDITION 
TO  FORESTED  AREAS  IN  EAST  AND  CENTRAL  AFRICA 

VI 

ITINERARY  AND  COMMENTS 


By  Arthur  Loveridge 


With  Four  Plates 


CAMBRIDGE,  MASS.,  U.S.A. 

PRINTED    FOR    THE    MUSEUM 

July,  1944 


No.  5.  —  Scientific  Results  of  a  Fourth  Expedition  to  Forested  Areas  in 

East  and  Central  Africa 

VI 

Itinerary  and  Comments 

By  Arthur  Loveridge 

INTRODUCTION 

For  various  reasons  it  would  appear  inopportune  at  the  present 
juncture  to  devote  the  time  necessary  to  elaborating  the  ecological  and 
zoogeographical  data  accumulating  from  this,  and  previous,  expedi- 
tions so  generously  sponsored  by  the  John  Simon  Guggenheim  Me- 
morial Foundation  of  New  York. 

On  several  occasions,  however,  correspondents  in  Africa  have 
written  requesting  specific  information  about,  or  the  latitude  and 
longitude  of,  some  of  the  more  obscure  localities  mentioned  in  the 
reports  already  printed,  so  that  it  would  seem  advisable  to  publish 
the  itinerary  without  further  delay.  A  synopsis  of  it  has,  indeed,  been 
furnished  in  the  caption  accompanying  Plate  1,  and  it  has  not  been 
thought  necessary  to  enlarge  on  this  in  respect  to  places  where  only 
a  single  night  was  spent,  with  consequently  little  collecting. 

For  the  others  I  am  now  supplying  the  latitude,  longitude,  altitude, 
the  all-important  meteorological  conditions  prevailing  at  the  time  of 
my  visit,  duration  of  stay,  and  position  of  camp.  This  in  turn  is  fol- 
lowed by  a  brief  survey  of  the  vegetational  environment,  or  a  reference 
to  where  such  an  account  has  been  published  elsewhere. 

The  composition  of  the  fauna  in  relation  to  life  zones  receives  atten- 
tion, more  particularly  for  localities  in  Tanganyika  Territory  where  it 
has  greater  significance  on  account  of  our  knowledge  being  more  com- 
plete. When  one  contemplates  the  vastness  of  the  largely  trackless 
forests  in  Uganda,  one  feels  that  the  results  accruing  from  a  stay  of 
two  weeks  in  areas  of  from  120  to  ISO  square  miles  in  extent,  is  alto- 
gether too  inadequate  to  justify  one  in  embarking  on  detailed  specu- 
lation as  to  what  does,  or  does  not,  occur. 

Species  of  exceptional  interest  receive  mention  where  it  is  thought 
that  they  may  assist  in  visualizing  the  environment,  or  where,  because 
of  their  rarity,  other  zoologists  visiting  the  locality  would  wish  to 
know  of  their  occurrence.  Special  reference  is  made  to  forms  of  which 
the  type  happens  to  have  come  from  the  particular  locality  under 
discussion. 


192 


bulletin:  museum  of  comparative  zoology 


The  groups  already  reported  upon  in  this  Bulletin  are  as  follows : 


Speci- 

New to 

Volume 

Pages 

Class 

mens 

Species 

M.C.Z. 

89 

145-214 

I  Mammals 

812 

116 

40 

89 

215-275 

II  Birds 

809 

240 

10 

91 

183-234 

III  Crustacea 

389 

20 

4 

91 

235-374 

IV  Reptiles 

1802 

151 

17 

91 

375-436 

V  Amphibians 

1081 

77 

10 

It  has  been  somewhat  of  a  disappointment  that  no  one  has  been  able 
to  undertake  the  identification  of,  or  to  report  upon,  the  fairly  exten- 
sive collections  of  other  invertebrate  groups  such  as  mollusks,  myria- 
pods,  and  earthworms.  In  addition  to  zoological  material,  we  pur- 
chased from  Baamba  and  Banyaruanda  tribes  1013  ethnological 
items,  with  a  gross  weight  of  almost  one  ton,  which  were  sent  by 
truck  and  train  over  a  thousand  miles  to  the  port  of  shipment. 

For  part  of  the  data  regarding  altitudes,  temperature,  composition 
of  forests,  etc.,  I  am  indebted  to  "Uganda"  by  Messrs  H.  B.  Thomas 
and  R.  Scott,  1935,  and  to  the  "Handbook  of  Tanganyika  Territory" 
by  G.  F.  Sayers,  1930;  both  mines  of  reliable  information. 

There  remains  only  to  again  express  my  deep  gratitude  to  the  John 
Simon  Guggenheim  Memorial  Foundation,  and  to  Dr.  Thomas  Bar- 
bour, Director  of  the  Museum  of  Comparative  Zoology,  for  making 
possible  this  expedition,  of  whose  harvest  of  herpetological  material  I 
am  now  able  to  take  full  advantage  in  the  revisionary  studies  of 
African  lizards  upon  which  I  am  engaged. 


ITINERARY 
UGANDA 

Mabira  Forest,  Kyagwe  (Chagwe).   0°24'  N.,  33°0'  E.    Alt.  4000  feet. 

As  the  rains  were  still  continuing  we  accepted  the  generous  offer  of 
Mr.  L.  Jarvis  to  occupy  the  late  Major  Cuthbert  Christy's  old  house 
at  Mubango,  a  rubber  and  coffee  plantation  in  the  southern  end  of 
the  forest,  five  miles  north  from  Najembe,  which  is  twenty  miles 
southeast  of  Jinja  on  the  Jinja-Kampala  road. 

Arrived  at  noon  on  November  5,  and  left  on  the  21st. 


loveridge:  African  itinerary  and  comments  193 

A  heavy  shower  amounting  to  .76  inches  fell  on  the  afternoon  of  our 
arrival,  .17  the  following  day,  and  .75  on  the  14th.  Actually  a  shower 
or  two  occurred  almost  daily  but  the  precipitation  was  too  small  to 
be  registered.  The  average  annual  rainfall  at  Mubango  is  55  inches. 
The  daily  temperature  at  7  a.m.  averaged  66°,  and  at  noon  80°. 

We  had  come  here  largely  because  Major  Christy's  labours  had  made 
the  Mabira,  which  covers  120  square  miles,  type  locality  for  Leptopelis 
n.  christyi,  Hylambates  verrucosus,  Miodon  g.  christyi,  and  Aparallactus 
christyi  (=  A.  modestus).  Of  these  we  were  successful  in  securing  only 
the  first  and  last. 

Despite  what  appeared  to  me  to  be  ideal  conditions,  I  personally 
captured  only  three  snakes  during  as  many  days  primarily  devoted  to 
searching  for  them  both  in  the  forest  and  along  its  edge,  where  drifts 
of  leaves  between  buttress  roots  and  rotting  logs  provided  suitable 
retreats.  On  four  succeeding  days  I  employed  an  allegedly  expert 
snake-catcher  who  got  nothing  during  that  entire  period.  Yet  some 
two-score  labourers,  engaged  in  clearing  undergrowth  on  the  planta- 
tion, brought  in  snakes  at  the  rate  of  nearly  two  a  day,  and  of  these 
the  first  eleven  snakes  represented  ten  different  species !  This  astonish- 
ing variety  continued  until  we  actually  had  eighteen  species  for  a  total 
of  twenty-seven  snakes.  Of  these  one — Dipsadoboa  unicolor — was  new 
for  Uganda,  another — Bothrophthalmus  I.  lineatus — constituted  the 
most  easterly  record  for  the  species,  though  its  presence  here  had  been 
forecast  by  Lt.  Col.  Pitman. 

Pitman  (1934,  Uganda  Journal,  1,  pp.  7-16)  has  furnished  an  inter- 
esting account  of  the  Mabira  and  its  fauna  and  flora,  rendering  un- 
necessary further  details  here.  Lizards,  with  the  exception  of  the 
arboreal  Algiroides  africanus,  were  disappointing,  being  of  widespread 
species.   A  single  chelonian  was  encountered. 

The  dense  undergrowth  rendered  bird  collecting  extremely  difficult 
and  only  twenty  skins  were  prepared  during  the  fortnight.  More  than 
a  score  of  mammals  were  taken,  however,  including  such  forest  forms 
as  Cercocebus,  Cercopithecus,  Hcliosciurus,  Tamiscus,  Protoxerus 
Cephalophus  and  an  arboreal  pangolin  which  was  described  as  a  new 
race  (Phataginus  tricuspis  mabirae). 

Budongo  Forest,  Bunyoro.     1°42'  N.,  31°24'  E.     Alt.  4000  feet. 

Camp  was  pitched  in  the  mahogany  nursery  at  BISU  beside  the 
Buchanan  Saw  Mills,  so  that  full  advantage  might  be  taken  of  the 
three-mile-long  corridor  cut  into  the  heart  of  the  forest. 

Arrived  in  afternoon  of  November  22,  and  left  on  December  7th. 

Heavy  downpours  occurred  at  intervals  during  our  stay  for  the  rain- 


194  bulletin:  museum  of  comparative  zoology 

fall  here  is  well  distributed  throughout  the  year,  the  annual  average 
being  65  inches.  During  our  visit  the  daily  temperature  at  7  a.m.  aver- 
aged 60°,  and  at  noon  82°. 

Situated  on  the  north-north-westerly  slope  above  the  escarpment 
towards  the  northern  end  of  Lake  Albert,  Budongo  is  rich  in  mahogany 
and  consequently  considered  the  most  valuable  of  all  Uganda's  lovely 
forests.  Ironwood  trees  are  also  plentiful,  in  some  areas  forming  over 
50%  of  the  canopy  and  being  largely  responsible  for  the  maintenance 
of  its  evergreen  appearance.  The  forest  survey  reveals  that  the  second 
story  stratum,  attaining  about  80  feet,  is  composed  in  part  of  such 
genera  as  Celtis  and  Funtumia  spp.,  while  the  third  story  is  scanty. 
Bordering  the  forest  and  at  the  edge  of  clearings,  a  dense,  often  im- 
penetrable, undergrowth  of  bush  has  sprung  up;  but  in  the  depths  of 
the  forest  where  the  canopy  was  contiguous  one  might  wander  at  will 
or  chase  the  frogs  which  went  leaping  over  the  damp  leaves  that  car- 
peted the  forest  floor. 

Our  visit  added  four  amphibia  to  the  Uganda  list,  the  most  in- 
teresting being  Rana  christyi,  for  Budongo,  covering  about  180  square 
miles,  shows  very  close  faunal  relationship  with  the  great  Ituri  Forest 
lying  away  to  the  west  in  the  Belgian  Congo. 

This  is  borne  out  by  a  number  of  interesting  herpetological  records 
recently  resulting  from  the  collections  made  here  by  Mr.  W.  J.  Eggel- 
ing,  Conservator  of  Forests,  whom  we  had  the  pleasure  of  meeting. 
This  West  African  complexion  was  reflected  by  7  of  the  9  species  of 
snakes,  and  4  of  the  7  kinds  of  lizards  collected,  though  one  of  these — 
Algircrides  africanus,  is  more  central. 

Budongo  is  type  locality  for  7  valid,  and  2  invalid,  races  of  birds, 
only  one  of  which  I  succeeded  in  getting,  this  was  the  gorgeous  little 
forest  kingfisher  {Myioceyx  lecontei  ugandae)  which  obligingly  alighted 
on  a  stump  in  camp. 

Of  the  10  species  of  mammals  secured,  only  4  can  be  considered 
definitely  western,  the  best  being  a  huge  horseshoe  bat  (Hipposideros 
cyclops)  netted  in  a  clearing  deep  in  the  forest,  where  was  also  the  great 
tree  on  which  I  shot  four  small  chipmunk-like  squirrels  (Tamiscus 
alexandri).  Here  too  we  trapped  a  strange  harsh-furred  mouse  (Lop- 
huromys  aquilis  subsp.)  whose  race  proved  unidentifiable;  perhaps  a 
series  might  reveal  it  as  an  undescribed  subspecies. 

Kibale  Forest,  Tow.     0°24'  N.,  30°24'  E.     Alt.  4200  feet. 

Camp  was  made  near  the  Duta  River  where  it  crosses  the  new  road 
being  cut  through  the  forest  about  20  miles  southeast  of  Fort  Portal. 

Arrived  late  on  December  8,  and  left  on  the  18th. 


loveridge:  African  itinerary  and  comments  195 

Thunderstorms,  accompanied  by  rain,  occurred  almost  every  after- 
noon, while  on  two  days  an  exceptionally  heavy  downpour  continued 
on  throughout  the  night.  The  annual  average  rainfall  is  56.87  inches. 
During  our  stay  the  daily  temperature  at  7  a.m.  averaged  59°,  and  at 
noon  77°. 

Kibale,  together  with  the  adjacent  Itwara  and  Muhangi  Forests, 
covers  about  276  square  miles.  The  northern  portion,  traversed  by 
the  Kampala-Fort  Portal  Road,  is  impenetrable  on  account  of  the 
dense  thicket  undergrowth.  Acting  on  the  advice  of  Mr.  W.  J.  Eggel- 
ing,  therefore,  we  selected  the  southern  portion  as  being  more  open, 
the  enormous  buttressed  trees  sufficiently  dense  as  to  prohibit  under- 
growth in  many  areas.  After  penetrating  the  forest  about  a  mile-and- 
a-half  Ave  had  to  camp  for  the  Dura  River  was  still  unbridged;  beyond 
the  river  the  road  continued  for  another  two-and-a-half  miles  through 
beautiful  forest,  and  it  was  along  this  stretch  that  most  of  my  collecting 
was  done. 

Excursions  along  this  road  during  and  immediately  after  heavy  rain 
were  productive  of  sundry  snails,  slugs  and  worms,  the  latter  very 
numerous  and  one  species  frequently  a  couple  of  feet  in  length.  This 
larger  species,  with  a  diameter  as  great  as  one's  thumb,  was  remarkably 
iridescent.  Beyond  the  forest  proper  these  worms  were  encountered 
in  muddy  soil  beside  a  stream  which  flowed  through  a  patch  of  palm 
forest  much  frequented  by  elephant.  Butterflies  in  bewildering  variety 
surpassed  anything  which  I  had  seen  anywhere  in  East  Africa. 

It  was  disappointing,  therefore,  to  find  the  herpetofauna  extremely 
scarce,  not  merely  in  species  but  in  individuals  also.  All  our  efforts  re- 
sulted in  obtaining  only  7  species  of  reptiles  and  7  of  amphibians, 
though  there  was  some  compensation  in  the  fact  that  all  were  exclu- 
sively sylvicoline  species  with  the  exception  of  Agama  atricollis  and 
Bujo  r.  regularis  which  are  equally  at  home  in  the  savanna.  The  head- 
man of  the  gang  engaged  in  felling  trees  with  which  to  bridge  the 
Dura,  informed  me  that  they  encountered  from  1  to  3  snakes  only  per 
month.  During  the  week  the  only  one  they  brought  me  was  a  Rham- 
nophis  a.  elgonensis.  It  was  this  paucity  of  poikilothermous  creatures 
that  decided  me  to  leave  after  ten  days  instead  of  remaining  three 
weeks  as  originally  planned. 

Monkeys  were  the  dominant  form  of  animal  life  and  no  fewer  than  5 
species,  of  which  I  collected  4,  were  observed  feeding  within  a  hundred 
yards  of  my  tent,  the  source  of  attraction  being  the  huge  spherical  fruit 
borne  by  a  certain  tree.  Of  a  dozen  species  of  mammals  collected,  a 
squirrel  (Funisciurus  p.  victoriae)  was  the  only  novelty. 


196  bulletin:  museum  of  comparative  zoology 

Bundibugyo,  Tow.    0°41'  N.,  29°56'  E.     Alt.  3200  feet. 

Stayed  at  the  comfortable  rest  camp  at  Saza  headquarters  for 
Bwamba  District. 

From  the  evening  of  December  19  to  26th. 

A  few  showers  occurred  during  the  week.  No  annual  rainfall  record 
was  available  as  the  area  has  only  been  opened  up  recently,  but  statistics 
are  being  collected  at  the  mission.  During  our  stay  the  daily  tempera- 
ture at  7  a.m.  averaged  55°,  and  at  noon  62°. 

This  area,  marked  on  the  Uganda  Survey  Map  of  1928  as  Bwamba 
Forest,  is  now  largely  covered  by  native  gardens,  including  plots  of 
coffee,  cotton,  and  bananas,  with  extensive  rice  cultivation  in  the 
swampy  areas.  Elsewhere  rank  grass  fifteen  feet  high,  scattered  trees, 
clumps  of  indigenous  oil  palms  (Elacis  guinecnsis)  and  patches  of 
forest.  The  latter,  said  to  amount  to  140  square  miles,  is  an  extension 
of  the  great  Ituri  Forest  on  the  opposite  bank  of  the  Semliki  River. 

Naturally,  therefore,  sylvicoline  snakes  are  plentiful  and  a  large  col- 
lection could  have  been  made  had  we  been  able  to  stay  longer.  At 
first,  however,  the  Baamba  mutilated  them  so  badly  that  many  had  to 
be  rejected  and  it  was  only  during  the  last  few  days  that  they  began 
to  arrive  in  good  condition.  At  least  14  of  the  16  species  secured  were 
forest  forms,  of  which  Boiga  puherulenta  and  Pscudohaje  goldii  were  the 
second  records  for  Uganda  and  Miodon  g.  collaris  the  first  for  this  race. 

Three  of  the  five  kinds  of  lizards  taken  were  widespread  savanna  or 
eastern  forest  forms.  No  attempt  was  made  to  collect  birds  and  the 
only  ones  preserved  were  the  swamp-loving  rail  (Sarothrura  p:  cen- 
tralis) and  warbler  (Prima  m.  immutabilis) . 

Except  for  a  young  duiker,  all  42  of  the  mammals  preserved  were 
rodents  representing  9  species  which  reflected  our  position  in  the  west- 
ern foothills  of  the  Ruwenzori  Mountains  overlooking  the  Semliki 
River,  some  being  montane  species,  others  races  of  savanna  rats. 

The  Baamba,  who  give  their  name  to  this  region,  are  a  forest  tribe 
of  semi-pigmy  stock  who  would  undoubtedly  prove  helpful  to  a  natural- 
ist working  in  this  country;  unfortunately,  as  already  indicated,  de- 
forestation has  resulted  in  encroachment  by  many  savanna  forms 
which  would  probably  preponderate  among  specimens  brought  in. 
The  big  tree  cobra  was  obtained  by  Bakonjo,  a  primitive  Bantu  people 
living  on  the  slopes  of  Ruwenzori.  Most  of  the  cultivation  was  being 
done  by  Batoro,  while  administration  was  largely  in  the  hands  of 
Bahima,  the  tall  Hamitics  and  former  overlords.  We  found  all  of  these 
people  most  friendly  and  only  too  eager  to  dispose  of  ethnological 
material,  much  of  it  superseded  by  changing  customs. 


loveridge:  African  itinerary  and  comments  197 

Bugoye,  Ruwenzori  Mountain*.  0°17'  N.,  30°13'  E.   Alt.  c.  5600  feet. 

( lamped  in  the  rest  camp  enclosure. 

December  26  to  28th,  1938,  and  January  21  to  24th,  1939. 

Occasional  showers.  We  had  left  our  thermometer  hanging  on  a  tree 
in  the  heart  of  Kibale  Forest  so  thereafter  we  were  unable  to  record 
temperatures. 

After  leaving  Bundibugyo  we  drove  over  the  northern  end  of  the 
Ruwenzori  Range,  which  is  about  30  miles  in  maximum  width,  through 
Fort  Portal,  skirted  the  eastern  flank  of  the  65-mile-long  range,  then 
turned  off  the  main  road  and  took  the  side  road  to  Bugoye  where  it 
terminates. 

Bugoye,  at  the  southeastern  foot  of  the  range,  is  not  only  the  type 
locality  of  Chamaeleo  h.  ellioti,  of  which  we  obtained  a  good  series,  but 
the  jumping-off  place  for  any  ascent  of  the  mountain  (16,800  feet) 
from  this  direction.  This  was  the  route  followed  by  the  British  Mu- 
seum Expedition  of  1908,  an  expedition  which  resulted  in  several  score 
of  new  vertebrates.  It  was  partly  in  the  hope  of  securing  topotypes, 
at  least  of  the  reptiles,  that  we  now  planned  to  visit  the  two  most 
readily  accessible  sites  occupied  by  Woosnam  and  his  party. 

We  stayed  at  Bugoye  only  as  long  as  it  was  necessary  to  get  to- 
gether sufficient  porters  for  the  ascent,  while  on  the  return  journey  I 
was  fully  occupied  with  packing  the  specimens  obtained  on  the  moun- 
tain before  the  lorry  should  arrive  to  pick  us  up. 

Consequently  little  collecting  was  done  in  this  uninteresting  spot 
whose  surrounding  slopes,  eroded  by  constant  rain,  are  clothed  only  in 
sparse  grass  with  thickets  and,  lower  down,  acacia  or  orchard  forest. 

Mubuku  (Mobuku)  Valley,  Ruwenzori.  0°24'  N.,  30°0'  E.  Alt.  6800  feet. 

Camped  in  deep  forest  between  the  Mubuku  and  Mahoma  Rivers, 
near  foot  of  Nyinabitaba  Ridge. 

Arrived  about  noon  on  December  29, 1938,  and  left  January  9,  1939. 

Rain  fell  on  every  day  except  one.  The  worst  aspect  of  the  situation 
was  the  absence  of  sunshine  due  to  overhanging  clouds  which  cast  an 
evening-like  gloom  and  stillness  over  the  forest  for  hours  on  end.  Sun- 
day was  the  best  day  with  7  hours  of  sunshine,  on  Monday  about  half 
that  amount,  the  rest  averaged  between  1  and  2  hours  only.  The  tem- 
perature at  midday  was  probably  from  55°  to  66°.  Under  such  condi- 
tions collecting  was  difficult,  birds  were  silent  except  during  the  brief 
hours  of  sunshine  when  they  were  engaged  in  feeding  on  the  forest 
canopy  well  out  of  range.  Drying  of  skins  and  other  specimens  became 
a  constant  nightmare. 

Despite  the  sodden  state  of  the  undergrowth  only  a  toad  and  3 


198  bulletin:  museum  of  comparative  zoology 

species  of  frogs  were  encountered,  all  the  latter  (Rana  f.  angolensis, 
Phrynobatrachus  graueri,  and  Hyperolius  ?  alticola)  previously  recorded 
from  the  mountain. 

Reptiles  were  limited  to  5  species  of  which  two  were  new  for  Ruwen- 
zori,  these  were  Cnemaspis  a.  elgonensis  and  Lygosoma  g.  graueri.  This 
little  pentadactyle  skink,  together  with  the  four-toed  L.  mcleagris,  of 
which  we  secured  topotypes,  were  relatively  plentiful.  Topotypes  were 
also  obtained  of  the  chameleons  (C.  j.  johnstoni  and  C.  xenorhinus) , 
the  latter  a  great  rarity  apparently  dwelling  in  the  forest  canopy. 
Naturally  C.  b.  rudis,  obtained  by  the  British  Museum  Expedition  at 
10,000  feet,  was  not  met  with  at  the  lower  level  but  I  was  surprised  to 
see  no  Lacerta  jacksoni,  a  species  that  has  been  taken  at  8500  feet. 

A  pair  of  crimson-winged  plantain-eaters,  presumably  Ruwenzoror- 
nis,  were  seen  near  camp  one  day  but  out  of  gunshot  in  the  tallest 
trees.  A  mountain  buzzard  (Buteo  oreophilus)  visited  our  lonely  camp 
and  fell  to  my  gun,  while  topotypes  of  half-a-dozen  passerines  were  also 
collected. 

Mammals  were  decidedly  scarce,  not  a  bat  was  seen,  and,  though  a 
net  was  spread  across  a  suitable  clearing,  nothing  was  taken.  Colobus 
were  seen  only  on  the  march  to  this  camp  and  all  the  monkeys  seen 
were  two  solitary  males  of  Cercopithecus  m.  stuhlmanni,  the  one  col- 
lected would  be  almost  a  topotype  of  the  synonym  carruthersi  which 
was  first  obtained  a  thousand  feet  higher  up  the  mountain.  During 
our  entire  stay  squirrels  (Heliosciurus  and  Tcnniscus)  were  observed 
four  times  only  .  Of  three  species  of  rodents  preserved  one  was  a  topo- 
type of  the  interesting  mouse  (Hylomyscus  d.  denniae).  A  red  duiker 
was  seen  once  and  heard  a  couple  of  times  but  attempts  to  collect  it 
failed.   Not  a  trace  of  any  carnivore  was  noted. 

Mihunga  Ridge,  Ruwenzori.     0°21'  N.,  30°3'  E.     Alt.  6000  feet. 

On  the  outward  journey  we  had  camped  for  the  night  beneath  the 
fig  tree — on  the  upper  Mihunga  Ridge  and  collected  a  few  specimens ; 
returning  we  passed  it  to  pitch  our  tents  on  the  lower  Mihunga  Ridge 
on  the  very  site  formerly  occupied  by  the  British  Museum  Expedition 
in  1908. 

Night  of  December  28,  1938.  Then  from  January  9  to  19th,  1939. 

We  had  fled  from  the  Mubuku  Valley  camp  on  account  of  adverse 
climatic  conditions,  now,  though  only  a  few  miles  distant  as  the  crow 
flies  we  seemed  to  be  in  another  world.  On  this  largely  treeless  spur, 
bounded  on  the  north  by  Weria  Ravine,  on  the  south  by  the  Kanyon- 
gorogoro  Ravine,  we  were  able  to  profit  by  the  hot  sunshine  which 
lasted  for  the  first  three  days  until  3  p.m.  and  was  then  followed  by 


loveridge:  African  itinerary  and  comments  199 

showers.  No  rain  at  all  occurred  on  the  four  days  following  while  dur- 
ing the  last  week,  though  the  sky  was  often  covered  by  fleecy  clouds 
through  which  the  sun  had  difficulty  in  breaking,  dark  clouds  formed 
only  in  the  late  afternoon  and  then  frequently  dispersed  without  preci- 
pitation in  our  vicinity,  though  at  times  rain  might  be  seen  falling 
elsewhere. 

Collecting  was  not  confined  to  the  6000-foot  ridge  for  excursions 
were  made  to  the  forested  heights  a  1000  feet  higher  as  well  as  to  the 
foot  of  the  ridge  where  it  tapered  out  into  the  swamps  of  the  Mubuku 
Valley.  For  descriptions  of  this  and  the  last  camp  the  reader  is  re- 
ferred to  Woosnam's  account  (1910,  Trans.  Zool.  Soc.  London,  19,  pp. 
5-24),  and  to  some  important  comments  by  Allen  and  Loveridge 
(1942,  Bull.  Mus.  Comp.  Zool.,  89,  p.  148). 

Almost  all  of  the  amphibians  (Bufo  r.  regularis,  Leptopclis  n.  christyi 
and  Phrynobatrachus  graueri)  and  most  of  the  9  species  of  reptiles 
taken,  came  from  the  swamp  about  500  feet  lower  than  our  camp. 
Here,  high  in  the  papyrus,  %were  coiled  the  green  vipers  (Atheris  n. 
nitschei)  topotypes  of  A.  woosnami  of  which  we  had  come  in  search. 
Here  also  we  obtained  the  first  Uganda  example  of  the  dwarf  chameleon 
(Brookesia  s.  boulengcri). 

Of  30  kinds  of  birds  collected,  4  were  topotypic,  while  each  morning 
our  traps  yielded  some  topotypic  rodent  until  we  had  10  of  the  species 
collected  at  Mihunga  by  Woosnam  and  his  associates;  3  others  were 
almost  topotypic  the  types  having  been  taken  at  higher  altitudes. 
This  region  appears  to  be  as  rich  in  mammals  as  it  is  poor  in  amphi- 
bians for,  in  addition  to  the  score  of  species  collected,  we  found  three 
sleeping  platforms  built  by  chimpanzees  in  a  tree  in  Weria  Ravine. 
The  Bakonjo  informed  me  that  these  platforms  are  made  by  itinerant 
chimpanzees  which  come  from  the  forests  of  the  Mubuku  in  search 
of  bananas  and,  when  benighted,  construct  a  platform.  We  saw  one 
such  individual  while  wTe  were  engaged  in  collecting  in  the  swamp. 

Nyakabande,  Kigezi.     1°44'  N.,  29°45'  E.     Alt.  6925  feet. 

Stayed  at  the  commodious  rest  camp. 

Arrived  at  noon  on  January  25  and  left  on  30th.  Returning  from 
Mushongero  on  February  4,  we  had  to  wait  for  a  lorry  until  the  8th. 

Though  there  was  relatively  little  sunshine,  the  weather  was 
generally  fine  except  for  a  few  heavy  showers. 

Nyakabande  is  situated  in  a  lava-strewn  plain,  much  of  which  is 
industriously  cultivated  by  the  Banyaruanda  who  gather  up  the  larger 
blocks  of  lava  and  pile  them  on  the  periphery  of  each  small  plot,  where 
they  serve  as  a  windbreak.    There  is  no  water  in  the  immediate 


200  bulletin:  museum  of  comparative  zoology 

vicinity,  and  perhaps  it  is  just  as  well  not  to  visit  the  foul  pool  from 
which  your  water  supply  is  likely  to  come.  Presumably  it  was  from 
this  pool  that  the  four  species  of  amphibia  obtained  here  came. 

Nyakabande  is  about  six  miles  from  Kisolo  (the  rendering  given  by 
the  Uganda  Survey  on  its  map  A  530  of  1928,  often  misspelled  Kisoro 
or  Kissolo),  type  locality  of  the  toad  I  named  Bufo  r.  kisoloensis, 
whose  validity  can  no  longer  be  maintained  as  a  result  of  the  fresh 
material  obtained  at  Nyakabande  and  Mushongero. 

The  reptile  fauna  of  these  two  localities  is  essentially  similar  as  we 
got  only  three  species  at  the  former  not  taken  also  at  the  latter,  which 
is  reached  by  a  very  arduous  climb  over  the  mountains. 

In  the  rest  house  grounds  we  found  a  weaver  (Ploceus  n.  graueri) 
nesting  in  a  vociferous  colony  of  the  superficially  similar  P.  c.feminina. 
At  Mushongero  we  obtained  a  topotype  wagtail.  (Motadlla  c.  wellsi) 
and  a  pair  of  flycatchers  (Alseonax  a.  ruandae)  which  race  was  de- 
scribed from  Bufundi  on  nearby  Lake  Bunyonyi. 

At  both  localities  the  Banyaruanda  apparently  eat  the  huge  mole 
rats  (Tachyoryctes  ruandae),  for  they  brought  in  as  many  as  I  would 
take.  The  species  was  first  collected  on  Mt.  Muhavura  which  is  in  full 
view  of  the  rest  house  at  Nyakabande.  More  valuable  was  an  otter 
(Lutra  m.  tenuis)  which  I  shot  in  the  lake  near  Mushongero,  for  it  was 
from  this  lake  that  Hinton  described  the  synonym  L.  m.  mutandae. 

The  main  object  of  our  stopover  at  Nyakabande,  however,  was  to 
purchase  ethnological  material  from  the  teeming  tribes  inhabiting  this 
upland  plain.  Collectively  known  as  Banyaruanda,  they  apparently 
consist  of  a  small  admixture  of  Hamitic  overlords,  the  cattle-owning 
Batusi  and  Bahororo.  The  bulk  of  the  population  consisting  of  Bantu 
agriculturists  known  as  Bahutu,  while  a  few  semi-pigmy  Batwa  are 
present,  chiefly  in  the  vicinity  of  the  lakes. 

Mushongero,  Lake  Mutanda,  Kigezi.  1°46'  N.,  29°41'  E.  Alt.  5925  feet. 

Stayed  at  the  rest  camp,  which  is  situated  on  a  little  peninsula  pro- 
jecting from  the  east  bank  near  its  northern  end. 

Arrived  late  in  the  afternoon  of  January  30,  and  left  early  on 
February  4th. 

Frequent  rainstorms  swept  across  the  lake  whose  mountain-girt 
northern  half  appeared  to  attract  and  hold  the  heavy  black  clouds 
which  overhung  it  during  much  of  our  brief  visit.  When  the  sun  was 
hidden  it  was  decidedly  chilly. 

Except  for  its  extensive  papyrus  swamps,  Lake  Mutanda  offers  little 
indication  of  its  proximity  to  the  equator.  In  fact  with  its  numerous 
tree-covered  islets  and  purplish  mountains  its  general  appearance  is 


loveridge:  African  itinerary  and  comments  201 

not  unlike  that  of  a  Scottish  loch.  The  brambles,  bracken,  and  short, 
wind-swept  grass  clothing  the  lower  slopes  of  the  mountains  all  suggest 
that  it  was  with  good  reason  that  these  uplands  have  been  called  the 
Switzerland  of  Central  Africa. 

As  for  the  name  Mushongero  (misspelt  Mushungero  on  all  my 
labels),  I  suspect  that  there  is  some  connection  between  it  and  Mus- 
hungwe,  the  Lugezi  name  for  leach,  for  these  loathsome  creatures 
swarm  in  the  shallow  waters  in  the  vicinity  of  the  rest  camp.  Crabs 
(Ngara:  Lugezi)  were  also  not  uncommon,  and  the  series  secured  have 
been  described  by  my  colleague,  Dr.  F.  A.  Chace  Jr.,  under  the  name 
of  Potamon  (Geothelphnsa)  mutandensis. 

The  waters  of  the  lake  were  teeming  with  Xenopus  1.  bunyoniensis, 
originally  described  from  nearby  Lake  Bunyonyi,  but  search  of  the 
papyrus  both  by  day  and  night  residted  only  in  the  capture  of  a  single 
example  of  the  genus  Hyperolius,  and  that  unidentifiable!  The  most 
perplexing  aspect  of  the  amphibian  fauna  was  the  intermediate  condi- 
tion of  many  Rana  fuseigula,  some,  as  one  would  expect  from  the  ter- 
rain, were  typical,  others  showed  the  longer  hind  limb  of  the  sylvicoline 
race  chapini.  Three  other  ranids  taken  here  also  seemed  to  suggest 
that  some  deforestation  had  taken  place. 

This  was  again  the  case  with  a  couple  of  the  dozen  species  of  rep- 
tiles collected.  \Ye  had  gone  to  Mushongero  primarily  in  search  of  a 
blind  snake  obtained  there  by  Col.  Pitman;  hoping  that  an  adequate 
series  of  the  creature  might  settle  its  uncertain  status.  In  this  we  were 
successful,  the  9  examples  undoubtedly  referable  to  Typhlops  blanfordii 
lestradci,  originally  described  as  a  fidl  species  from  nearby  Ruhengeri 
but  obviously  related  to  blanfordii  of  the  Ethiopian  highlands;  another 
link  with  the  latter  region  is  the  presence  at  Mushongero  of  the  little 
slug-eater  (Duberria  I.  abyssinica). 


BELGIAN  RUANDA 

Kiraga  near  Kisenyi,  Lake  Kivu.  2°38'  S.,  29°1S'  E.   Alt.  c.  5800  feet. 

Camp  was  pitched  in  a  plantation  of  eucalyptus  through  which  the 
road  passes  about  three  miles  above  Kisenyi,  a  township  situated  on 
the  shores  of  Lake  Kivu  at  4800  feet. 

Arrived  in  afternoon  of  February  8,  and  left  early  on  the  13th. 

Some  showers. 

Most  of  our  collecting  at  Kiraga  was  done  along  the  banks  of,  and 
in  the  ravine  cut  by,  the  Kisenyi  River,  which  cascaded  over  falls  just 


202  bulletin:  museum  of  comparative  zoology 

below  our  camp.  The  ravine  was  more  or  less  choked  by  luxuriant 
growths  of  grass  and  sedge,  but  in  patches  cleared  for  gardens  we 
turned  over  piles  of  vegetable  debris  and  sought  our  quarry  in  the  ex- 
tensive plantations  of  bananas. 

Only  4  species  of  amphibia  and  6  of  reptiles  were  taken,  none  of 
especial  interest  with  the  possible  exception  of  Lacerta  jacksoni  which, 
in  the  absence  of  trees,  has  adapted  itself  to  a  terrestrial  life.  By  far 
the  most  important  of  half-a-dozen  mammalian  species  were  a  pair  of 
skunk-like  zorillas  (Poecilogale  a.  doggetti). 

As  we  were  debarred  from  killing  anything  but  "vermin"  on  ac- 
count of  our  permit  for  scientific  collecting  not  having  arrived  from 
Stanleyville,  we  caught  the  first  boat  from  Goma  (Ngoma),  type 
locality  for  a  burrowing  viper  (Atractaspis  schoutedeni),  which  appears 
to  be  based  on  a  slightly  aberrant  example  of  the  widespread  A.  irre- 
gularis, which  we  got. 


BELGIAN  CONGO 

Mamm  Bay,  Idjivi  Island.     2°12'  S.,  29°0'  E.     Alt.  4788  feet. 

Camped  on  the  lawn  of  Mons.  van  der  Berck  v.  Heemstede's  estate. 

Arrived  at  dusk  on  February  14  and  left  on  16th,  returning  March 
6th. 

Our  arrival  was  greeted  by  torrents  of  rain  and  waves  lashed  by  a 
gale. 

Wading  in  thigh  boots  among  the  sedges  of  the  Bay,  and  aided  by  a 
flashlight,  I  was  able  to  capture  topotypes  of  Ilyperolius  kivuensis, 
Icwidjwicnsis,  kandti,  and  macrodactylus,  and  show  that  actually  only 
two  sexually  dichromatic  species  are  present.  The  types  had  been 
collected  by  the  poet  Kandt,  who  had  made  his  home  on  Idjwi  Island. 

Upper  Mulinga  River,  Idjwi  Island,  c.  2°8'  S.,  29°3'  E.  Alt  ^500  feet. 

Camped  beside  a  footpath  where  it  crosses  a  stream  known  as  the 
Upper  Mulinga.  This,  I  imagine,  was  about  900  to  1000  feet  below 
the  800  metre  summit  of  the  mountain,  which  dominates  the  island. 

Arrived  on  February  16  and  left  at  noon  on  March  6th. 

The  weather  was  very  varied,  we  enjoyed  much  sunshine  when  it 
would  be  quite  hot;  violent  thunderstorms  accompanied  by  lashing 
rain  were  not  uncommon,  however,  and  always  resulted  in  a  consider- 
able drop  in  temperature. 

1  Not  4500  feet  as  printed  on  p.  149  of  the  report  of  Mammals,  1942,  Bull.  Mns.  Comp.  Zool., 
89,  pj>.   147,  214.    I  am  indebted  to  Dr.  J.  P.  Chapin  for  pointing  out  this  error. 


loveridge:  African  itinerary  and  comments  203 

Day  after  day  I  made  excursions  up  the  mountain  to  the  extensive 
remnants  of  magnificent  forest,  even  then  being  destroyed  by  natives 
contrary  to  regulations;  policing  of  such  remote  spots  being  difficult. 
Apart  from  some  small  patches  of  forest,  our  immediate  vicinity  con- 
sisted of  pasture  land,  millet  fields,  dense  patches  of  sedge,  and 
swampy  areas  through  which  meandered  small  rivulets. 

It  was  in  this  latter  habitat  that  I  collected  most  of  the  8  species 
of  amphibia  taken,  but  the  choicest  of  all,  a  tiny,  long-fingered  male 
Arthrolcptis  xenochirus,  was  brought  to  me  by  a  native  lad.  Only  2  of 
the  species  were  savanna  forms. 

Of  the  25  species  of  reptiles  collected,  all  but  a  fourth  were  of  forest 
association  and  included  such  rarities  as  Miodon  g.  graueri  and  Algi- 
roides  vaucreselli.  The  island,  variously  spelt  Idschwi,  Kwidjwi,  and 
Kidjwi  by  the  Germans,  is  type  locality  for  Mabuya  m.  kicidjwicnsis 
and  Lygosoma  blochmanni.  Over  fifty  of  each  were  preserved,  suffi- 
cient to  demonstrate  that  the  former  does  not  differ  structurally  from 
true  M.  m.  maculilabris,  and  that  the  latter  is  constantly  three-toed. 
The  local  chameleon  appeared  to  differ  sufficiently  from  its  continental 
congeners  to  be  named  C.  d.  idjwiensis. 

Also  new  were  8  examples  of  a  warbler  (Apalis  eidos),  and  we  secured 
topotypes  of  Barbatula  kandti  =  Pogoniulus  b.  jacksoni,  Colivs  s. 
kiwuensis  and  Spinus  c.  frontalis;  in  all  42  species  of  birds  were  col- 
lected at  this  camp. 

Two  new  races  of  rodents  (Thamnomys  v.  kimiensis  and  Leggada  b. 
ablutus)  were  discovered,  the  former  just  behind  my  tent,  the  latter 
differing  from  the  typical  race  occurring  on  the  Ruwenzori  Moun- 
tains. An  interesting  aspect  of  the  mammalian  fauna  of  this  mountain 
on  Idjwi  was  that  a  third  of  the  21  species  collected  were  referable  to 
races  originally  described  from  Ruwenzori;  two  others  (Cercopithecus 
m.  schoutedeni  and  Lophuromys  a.  laticeps)  were  topotypes  of  local 
forms. 

TANGANYIKA  TERRITORY 

Ujiji,  Kigoma  District.     4°55'  S.,  29°42'  E.     Alt.  2800  feet. 

Camped  beneath  the  giant  mangoes  on  the  western  fringe  of  the 
town. 

Arrived  at  noon  on  March  9,  and  left  early  on  March  10th. 

Occasional  heavy  showers  following  periods  of  sultry  weather  and 
overcast  skies. 

As,  from  May  22  to  29,  1930,  I  had  already  visited  this  famous  old 


204  bulletin:  museum  of  comparative  zoology 

Arab  settlement  on  the  shores  of  Lake  Tanganyika,  a  description  of 
its  features  will  be  found  in  1933,  Bull.  Mus.  Comp.  Zool.,  75,  p.  23. 

The  purpose  of  my  visit  was  the  same  in  both  instances,  viz.  to  ob- 
tain examples  of  the  rare  Amphisbaena  phylofiniens,  known  only  from 
the  two  cotypes  described  in  1905.  On  the  first  occasion  I  was  unsuc- 
cessful but  learned  from  the  natives  that  it  was  to  be  found  in  Ruanda, 
a  region  of  rice  swamps  bordering  the  Luiche  River.  Almost  daily, 
therefore,  accompanied  by  two  assistants,  I  tramped  over  to  Ruanda, 
pausing  en  route  to  inform  each  passer-by  of  our  object.  Not  an  am- 
phisbaenid  did  we  find,  but  as  a  result  of  our  ceaseless  talk  4  speci- 
mens were  brought  in  by  natives.  Since  collecting  all  7  East  African 
members  of  the  family,  I  have  come  to  the  conclusion  that  the  habitat 
which  these  wormlike  creatures  require  is  one  of  moist  sand  or  laterite 
soil.  There  is  no  evidence  which  would  justify  the  view  that  they  are 
survivals  of  a  former  forest  fauna  in  this  region. 

While  searching  for  amphisbaenids  we  secured  a  good  series  of  a 
Congolese  skink  (Scelotes  t.  hemptinnei)  which  had  not  previously  been 
taken  in  Tanganyika.  Another  first  record  for  the  Territory  was  a 
"two-headed  snake"  {Chilorhinophis  gerardi)  though  I  had  postulated 
its  occurrence  at  the  southeast  end  of  the  lake  in  1933.  The  Angola 
race  ornatum  of  Lycophidion  capense  was  a  further  addition  to  the  fauna 
and,  from  a  distributional  point  of  view,  an  Aparallactus  c.  capensis  is 
interesting. 

Additions  to  the  fauna  among  the  9  species  of  amphibia  taken,  were 
Rana  m.  venusta  and  Hypcrolius  kivuensis;  topotypes  of  H.  udjijiensis 
and  argentovittis  were  also  captured. 

Kitaya,  Southern  Province.    10°40'  S.,  40°11'  E.   Alt.  300  feet. 

Tents  were  pitched  on  the  rest  camp  site  on  the  bank  of  the  Rovuma. 

Arrived  at  noon  on  March  24th  and  left  early  on  April  7th. 

Torrential  rains  fell  during  our  stay,  necessitating  the  employment 
of  porters  for  the  first  nine  miles  of  the  return  journey  to  Mikindani, 
i.e.  until  past  a  depression  of  water-logged  black  cotton  soil. 

Kitaya  is  a  village,  and  Liwale  headquarters,  on  the  north  bank  of 
the  Rovuma,  (Rowuma;  Ruvuma)  River,  fifteen  miles  inland  from 
Rovuma  Bay.  The  River  forming  the  southeastern  boundary  between 
Tanganyika  Territory  and  Mozambique.  Livingstone  safaried  there 
from  Mikindani  in  1866;  the  two  places  are  now  connected  by  a  very 
rough  motor  track  of  about  36  miles.  The  vegetation  is  typical  of  the 
coastal  belt;  baobabs,  orchard  forest,  and  rank  grass  on  the  higher 
ground,  which  is  very  sandy;  rice  fields  and  sedge  (Setaria  palmifolia) 
fringed  swamps  on  the  lower. 


loveridge:  African  itinerary  and  comments  205 

Huge  leeches  are  present  in  the  swamps,  ticks  in  the  long  grass. 
Aedes,  Glossina,  Haematopota,  Stomoxys,  Tabanus,  and  many  other 
biting  flies  were  an  ever-present  nuisance. 

The  purpose  of  our  visit  was  to  secure  topotypical  material  of  three 
species  of  sedge  frogs  {Hyperolius  eitrinus,  microps,  and  Megalixalus 
fiavomaculahis)  collected  by  Kirk  when  he  accompanied  Livingstone. 
In  this  we  were  eminently  successful  and  able  to  prove  that  the  last 
named  is  really  a  Hyperolius,  having  a  horizontal  pupil  in  life.  In 
addition  a  series  of  an  undescribed  race  of  this  genus  (//.  p.  rovumae) 
were  collected  and  described. 

We  had  to  depend  Aery  largely  on  our  own  efforts,  for  the  local 
people — Konde  and  Yao — were  strangely  disinterested  in  bringing  in 
reptiles  or  mammals,  in  fact  supplied  only  two  of  the  latter.  Crocodiles 
as  man-eaters  play  a  prominent  role  in  local  village  life.  Soft-shelled 
turtles  (Cycloderma  frenulum)  were  laying  and  in  consequence  may 
have  been  more  conspicuous  than  would  normally  be  the  case.  The  13 
species  of  snakes  might  almost  have  been  selected  as  representative  of 
the  most  widespread  African  forms!  Except  for  Ichnotropis  squamuhsa 
the  dozen  species  of  lizards  were  likewise  of  little  interest. 

Birdlife  was  wonderful,  being  particularly  rich  in  non-passerine 
species  such  as  parrots,  hornbills,  wood  hoopoes,  cuckoos,  and  wood- 
peckers. In  all  61  species  of  birds  and  a  dozen  different  kinds  of  mam- 
mals were  collected  during  the  13  days  spent  here.  Of  the  latter  a 
molossid  bat  (Mops  angolensis  orientis)  was  described  as  new. 

Mikindani,  Southern  Province.    10°17'  S.,  40°7'  E.    Alt.  20  feet. 

Tents  were  pitched  about  two  miles  north  of  the  centre  of  the  town- 
ship, which  gives  its  name  to  Mikindani  Bay  on  the  southeast  coast  of 
Tanganvika  Territorv,  on  a  little  rise  to  the  left  of  the  mainroad  to 
Lindi  after  one  passes  the  Government  pumping  station.  Another 
noisy  pumping  station  engine  has  given  the  name  of  Mchuchu  to 
this  area. 

First  landed  at  Mikindani  on  the  night  of  March  22  and  accom- 
plished some  collecting  on  the  following  evening  before  leaving  for 
Kitaya.  Returning  from  the  latter  place  on  April  7  in  torrential  rain, 
we  just  managed  to  get  the  tents  pitched  before  darkness  fell  on  the 
most  cheerless  conditions  of  the  whole  trip.  Left  by  dhow  for  Lindi, 
the  road  being  under  water  for  miles,  on  April  24th. 

Fully  0  inches  of  rain  fell  during  our  sixteen  days  stay,  on  some  days 
continuing  from  dawn  till  dusk.  The  greatest  single  precipitation 
recorded  was  1  3/16th  inches  on  April  10,  for  this  information  and 
other  kindnesses  I  am  indebted  to  Mr.  E.  A.  Leakey,  District  Officer. 
The  average  annual  rainfall  is  36  inches. 


206  bulletin:  museum  of  comparative  zoology 

Such  conditions  naturally  retarded  outdoor  studies  to  some  extent. 
Most  of  our  collecting  was  carried  out  in  the  eastern  environs  of  the 
township  and  relatively  little  in  the  immediate  vicinity  of  camp. 
Though  the  low  ground  surrounding  the  latter  was  largely  under  water, 
the  rainy  season  was  already  so  well  advanced  that  spawning  was  over 
for  the  majority  of  frogs,  which  were  already  widely  dispersed.  Of 
the  13  species  collected  only  one  (Arthrolcptis  xenodactylus)  has  any 
claim  to  forest  associations. 

Such  primeval  forest  as  may  have  been  here  long  since  disappeared, 
but  the  numerous  mango  and  other  trees,  to  say  nothing  of  baobab  and 
coconut,  result  in*  providing  conditions  acceptable  to  some  sylvicoline 
species.  Of  the  31  species  of  reptiles  taken,  however,  only  a  cobra 
(Naja  melanoleuca),  is  a  western  forest  form,  the  7'  6"  specimen  pro- 
viding the  most  southeasterly  record  for  the  species.  Topotypes  were 
collected  of  the  only  two  lizards  (Lygodactylus  g.  grotci  and  Amphis- 
baena  orientalis)  described  from  Mikindani. 

Of  the  43  species  of  birds  collected,  topotypes  were  shot  of  Francoli- 
nus  h.  grotci,  Lagnosticta  r.  rcichcnowi  =  haematocephala,  and  Uraegin- 
thus  b.  mikindaniensis  =  niassensis,  while  of  3  other  races  described 
from  here  we  had  obtained  specimens  from  nearby  Kitaya  and  Nch- 
ingidi. 

Only  7  kinds  of  mammals  were  obtained,  all  typical  of  the  coastal 
zone,  the  bat  Triaenops  afer  being,  perhaps,  the  most  interesting. 

Mbanja,  Southern  Province.  9°24'  S.,  39°45'  E.  Alt.  Sea  level  to  400 
feet. 

Camp  was  made  at  the  edge  of  Mitonga  (Metonge)  landing  field 
('aerodrome'),  circa  375  feet. 

Arrived  at  noon,  by  truck  from  Lindi,  on  April  25,  and  left  at  8  a.m. 
on  May  6,  1939. 

Heavy  showers  occurred  during  the  first  week;  the  second  was  prac- 
tically rainless,  the  heat  tempered  to  some  extent  by  the  southeast 
monsoon,  until  we  were  struck  by  a  gale  of  wind  and  rain  on  our  last 
night. 

Mbanja  (Mbanya),  about  10  miles  north  of  Lindi,  is  a  small  village 
situated  on  a  tidal  estuary  and  almost  surrounded  by  mangrove 
swamps.  The  chief  and  his  people  were  most  friendly  and  helpful. 
While  the  village  itself  is  on  clays  and  coral  rock,  the  valley  at  whose 
mouth  it  lies  is  largely  composed  of  black  cotton  soil  and  rich  mud  ex- 
tensively cultivated  by  the  industrious  inhabitants.  Their  principal 
products  being  ground  nuts,  potatoes,  and  mahoga,  with  a  few  paupau 
trees  and  coconut  palms  about  each  hut. 


loveridge:  African  itinerary  and  comments  207 

My  camp,  on  higher  ground,  was  largely  surrounded  by  orchard 
forest  heavily  interspersed  with  mango  trees  and  waist-high  grass 
whose  barbed  black  seeds  cause  considerable  discomfort  by  working 
through  clothing  and  even  puttees.  To  the  west  lay  dense  scrub  in- 
habited by  very  wary  squirrels  and  blue  monkey,  for  the  cry  of  the 
latter  was  heard  towards  sunset  on  several  occasions.  To  the  east, 
i.e.  between  camp  and  the  coast,  were  native  gardens  and  the  exten- 
sive Kikwetu  Sisal  Estate.  All  this  country  is  composed  of  bright  red 
and  very  porous  soil  derived  from  eroded  limestone. 

It  is  in  this  soil  that  Amphisbaena  ewerbecki,  described  from  here  by 
Werner  in  1910,  occurs.  Our  object  in  coming  to  Mbanja  was  solely 
to  get  a  series  and  in  this  we  were  entirely  successful.  The  soil  which 
furnished  a  congenial  habitat  for  the  amphisbaenid  was  favoured  by 
six  species  of  fossorial  snakes  which  we  collected  in  addition  to  9  other 
kinds  of  ophidia,  a  dozen  lizard  forms,  and  36  of  birds. 

Game  was  allegedly  shot  out  by  the  Germans,  who  maintained  a 
big  camp  near  here  during  1914-1917,  but  wild  pigs  were  common.  At 
night  a  solitary  little  antelope  emerged  from  the  scrub  to  feed  on  the 
foliage  of  the  ground  nuts.  Other  animals  seen  or  heard,  but  not  col- 
lected, were  red  elephant  shrews,  galagos,  baboons,  hares,  ratel  and 
jackal.  Man-eating  lions  had  been  causing  numerous  deaths  quite 
recently  in  neighbouring  villages. 

Rondo  Forest,  Southern  Province.   10°8'  S.,  39°12'  E.    Alt.  2700  feet. 

Camped  at  Nchingidi,  the  name  given  to  a  clearing  at  the  forest 
edge  approximately  three  miles  from  the  nearest  scattered  native  huts. 

Arrived  just  before  noon  on  May  9,  and  remained  until  the  21st. 

Each  evening  at  varying  times  from  sunset  (6  p.m.)  till  11  p.m.,  a 
succession  of  mist  clouds  blew  in  from  the  ocean  and  up  the  face  of  the 
escarpment  (at  whose  edge  my  tent  was  pitched)  to  condense  on  grass 
and  trees.  The  latter  literally  'rained'  upon  my  tent  with  every  gust 
of  wind.  At  daybreak  these  mist  clouds  hemmed  us  in,  completely 
shutting  out  the  view  of  the  opposite  escarpment,  invariably  persist- 
ing till  8  a.m.,  more  usually  9  a.m.,  occasionally  even  until  noon.  In 
addition  about  4  or  5  p.m.  there  were  sharp  showers,  while  sudden  and 
unexpected  storms  of  rain  swept  the  plateau  at  uncertain  intervals, 
and  frequently  heavy  downpours  occurred  at  night.  The  nights  were 
always  cool  but  when  the  sun  did  break  through  by  day  it  was  very  hot. 

Unfortunately  an  error  deprived  me  of  my  quinine,  and  repeated 
drenchings  when  far  from  camp  brought  on  a  fever  which,  so  far  as  I 
was  concerned,  halted  collecting  during  the  last  week. 

This  open  forest  is  situated  on  the  waterless  Rondo  Plateau,  about 


208  bulletin:  museum  of  comparative  zoology 

25  miles  sothwest  of  Lake  Rutamba,  (Lutamba),  itself  some  25  miles 
by  road  slightly  southwest  of  Lindi.  The  curious  thing  about  this 
forest,  of  which  mvidi  is  the  most  important  constituent,  is  the  entire 
absence  of  standing  water.  Even  the  heaviest  downpours  immediately 
disappeared  in  the  porous  sandy  soil.  To  obtain  water  for  domestic 
purposes,  the  local  natives  (Mwera)  made  a  daily  journey  of  three 
hours  (there  and  back)  to  the  foot  of  the  escarpment.  It  is  not  alto- 
gether surprising  that  with  water  so  scarce  these  people  should  con- 
sider washing  superfluous.  Though  very  friendly,  these  Wamwera 
only  began  to  busy  themselves  with  bringing  in  specimens  a  day  or 
two  before  our  departure. 

Despite  the  absence  of  water,  5  species  of  amphibia,  all  previously 
taken  on  the  Uluguru  Mountains  nearly  300  miles  to  the  north,  were 
collected.  Of  these  a  tiny  toad  (Bufo  micranotis  rondoensis)  differed 
sufficiently  to  warrant  description. 

This  isolated  plateau  had  produced  3  species  of  fossorial  reptiles 
(Amphisbaena  rondoensis,  Melanoseps  a.  rondoensis,  Typhlops  t.  ron- 
doensis) which  it  was  necessary  to  describe  as  new.  Here  also  we 
found  Chlorophis  maerops,  Bitis  gabonica,  and  Brookesia  brevicaudata 
which  occur  on  the  Usambara  Mountains  350  miles  to  the  north ;  even 
more  surprising  was  an  undoubted  Aparallactus  jacksonii  of  Kiliman- 
jaro. The  remaining  reptiles — 12  kinds  of  snakes  and  10  of  lizards 
were  representative  of  the  coastal  plain  herpetofauna. 

In  the  past,  principally  during  the  Great  War,  the  forest  suffered 
heavily  from. native  incursions.  Subsequently  these  refugees  were  re- 
moved but  evidence  of  their  destructive  occupation  were  to  be  seen 
in  the  numerous  clearings,  some  so  eroded  as  to  be  semiarid  areas  with 
only  the  scantiest  covering  of  grass,  others  grass-  or  bush-grown  and 
inhabited  by  such  non-sylvicoline,  house-dwelling  lizards  as  Hemi- 
dactylus  mabouia  and  Mabiiya  striata,  which  I  assume  to  have  been 
imported  by  human  agency. 

Bird  life,  though  not  abundant,  presumably  owing  to  the  absence 
of  standing  water,  held  promise  of  being  interesting  so  that  it  was  a 
disappointment  to  go  down  with  fever  just  as  I  was  turning  my  atten- 
tion to  the  avifauna.  A  beautiful  roller  {Eurystomus  glaucurus)  of 
Madagascar  was  preserved,  while  the  records  of  the  green -headed 
Oriole  (0.  chlorocephaius)  from  Mt.  Chiradzulu,  Nyasaland  and  the 
Usambara  Mountains  are  at  last  bridged  by  its  occurrence  on  the 
Rondo  Plateau.  Two  rare  flycatchers  (Batis  reichenoioi  and  Erythro- 
cercus  I.  thomsoni),  originally  described  from  Mikindani  and  the 
Rovuma  River  respectively,  were  also  among  the  25  species  obtained 
at  Nchingidi. 


loveridge:  African  itinerary  and  comments  209 

Doubtless  the  water  supply  is  also  the  reason  for  the  scarcity  of 
mammals,  of  which  only  9  species  were  taken,  the  most  interesting 
being  Rhynchocyon  p.  melanurus,  originally  described  from  Lindi.  I 
heard,  but  never  saw,  blue  monkey,  bushbuck,  and  a  small  duiker 
which  the  natives  called  naunde. 

Lindi,  Southern  Province.    10°0'  S.,  39°14'  E.    Alt.  50  feet. 

In  Lindi  Hospital  (one  week)  and  at  the  Beach  Hotel  (one  week). 

Arrived  on  May  22  and  sailed  on  June  5th. 

The  rains,  which  average  34  inches  per  annum,  were  over,  and  the 
clay  soil  already  baked  hard  by  a  tropical  sun. 

Lindi,  headquarters  for  the  Province  of  the  same  name,  is  situated 
in  Lindi  Bay  between  Kilwa  and  Mikindani.  A  population  of  nearly 
4000,  and  the  excellent  hygenic  conditions  of  the  township,  rendered 
collecting  difficult  during  the  week  that  I  daily  awaited  the  arrival  of 
a  steamer.  All  the  same  we  preserved  15  species  of  amphibia  and 
reptiles,  16  of  birds,  and  5  of  mammals. 

On  our  very  first  hunt,  however,  we  uncovered  the  oriental  blind 
snake  (Typhlops  braminus),  the  fourth  and  fifth  examples  to  be  taken 
in  the  Territory.  T.  s.  mucruso  was  the  only  other  snake  obtained 
though  bundles  of  thatching  grass,  piles  of  palm  fronds,  and  heaps  of 
rubbish,  were  turned  over  a  wide  area.  Amphisbaena  cwcrbccki  was 
the  only  lizard  of  interest.  An  intelligent  fisherman  told  me  that  four 
turtles  are  occasionally  encountered  in  the  Bay,  he  described  the  luth, 
loggerhead,  hawksbill,  and  green  turtle.     , 

Siga  Caves,  Tonga  Province.   5°6'  S.,  29°4'  E.   Alt.  c.  150  feet. 

Camp  was  pitched  on  the  trail  leading  to  the  caves  but  about  a  mile 
from  the  entrance. 

Arrived  on  June  7  and  remained  until  the  17th. 

A  few  heavy  rainstorms  occurred  at  infrequent  intervals,  the  rainy 
season  being  over. 

The  Siga  Caves,  I  took  the  name  from  the  government  signpost 
erected  at  the  turn  off  from  the  main  Tanga  to  Mombasa  road,  have 
nothing  to  do  with  the  Sigi  River  a  few  miles  to  the  north.  They  are 
sometimes  called  the  Amboni  or  Mkulumusi  Caves  for  the  main  en- 
trances are  less  than  fifty  yards  from  the  crocodile-infested  Mkulumusi 
River.  The  numerous  caves  are  waterworn  and  in  past  times  un- 
doubtedly formed  an  underground  channel  for  the  river.  There  are 
dubious  stories  of  the  caves  extending  for  a  mile,  but  at  the  time  of  our 
visit  every  passage  was  flooded  to  within  a  hundred  yards  of  the 
entrance. 

The  fifty  acres  of  forest  which  surrounds  them  has  suffered  con- 


210  bulletin:  museum  of  comparative  zoology 

siderably,  in  fact  it  seemed  to  me  that  little  remains  but  a  scattering 
of  fine  trees  surrounded  by  secondary  growth  and  much  scrub.  I 
might  add  that  the  caves  are  held  in  superstitious  veneration  by  the 
natives,  and  propitiatory  rites,  of  which  I  have  given  a  brief  account 
(1940,  Scientific  Monthly,  51,  pp.  22-35)  were  held  there  at  the  time 
of  our  visit.  I  imagined  that  the  presence  of  this  "spirit"  was  the 
reason  for  the  absence  of  native  squatters :  later  I  learned  that  during, 
and  immediately  following,  the  Great  War,  natives  had  moved  in  but 
were  turned  out  again  by  the  Administration  on  account  of  their  reck- 
less destruction  of  trees. 

All  11  species  of  amphibia,  as  well  as  13  kinds  of  reptiles  obtained  at 
Siga,  were  typically  coastal  plain.  One  gecko  (Cncmaspis  a.  africanus) 
was  definitely  sylvicoline,  while  five  others  proved  of  assistance  in  de- 
fining a  new  coastal  race  named  Hcmidactylus  t.  barbouri. 

No  serious  attempt  was  made  to  collect  birds  in  so  well-worked  a 
region,  and  only  8  species  were  shot,  the  choicest  being  a  pair  of 
migrant  Malagasy  egrets  (Egretta  g.  dimorpha)  and  my  first  bat-eating 
hawk  (Machaerhamphus  a.  anderssoni). 

The  latter  was,  of  course,  attracted  by  the  thousands  of  bats  which 
made  the  caves  their  home.  Of  these  bats,  96,  representing  4  species, 
were  collected ;  they  ranged  in  size  from  the  huge  Hipposideros  g.  gigas 
to  the  modest  Miniopterus  minor.  Galagos  (G.  c.  lasiotis)  were  also 
plentiful,  their  strange  cries,  mingling  with  the  staccato  bark  of  hyrax 
and  the  hooting  or  screeching  of  owls,  rendered  our  nights  the  noisiest 
of  the  entire  safari.  The  10  species  of  mammals,  all  troglodyte  or 
arboreal  except  for  a  spiny  mouse  {Acomys  w.  wilsoni),  were  of  coastal 
rather  than  of  forest  affinities. 

Amboni  Estate,  Tanga  Province.   5°3'  S.,  39°3'  E.   Alt.  300  feet. 

Our  tents  were  pitched  beside  a  small  area  of  forest,  largely  second- 
ary, which  is  being  carefully  preserved  by  the  Estate  Management. 
As  Amboni  Estate  covers  nearly  80  square  miles,  the  precise  location 
may  be  somewhat  important,  we  were  located  about  a  mile  above 
Mabokweni  Village  and  surrounded  on  three  sides  by  sisal  planta- 
tions. 

Arrived  on  June  17  and  remained  until  the  27th. 

Rain  occurred  in  scattered  showers  on  about  four  cloudy  days. 

It  seemed  strange  that  frogs  should  be  assembling  at  the  conclusion 
of  the  rainy  season,  yet  a  visit  to  a  rice  swamp  close  to  Mabokweni 
Village  on  the  evening  of  our  arrival,  resulted  in  the  capture  of  154 
polypedatid  frogs  of  9  different  species,  the  females  of  Hylambates 
maculatus  certainly  were  gravid  and  about  to  spawn. 


loveridge:  African  itinerary  and  comments  211 

Between  this  swamp  and  our  camp  was  an  extensive  area  that  had 
been  under  sisal  since  1913.  It  had  recently  been  cleared  and  tractors 
were  now  engaged  in  spreading  the  vegetable  debris  which  had  been 
piled  in  long  rows  and  left  to  rot  for  two  months.  By  following  the 
twelve-ton  tractors  to  and  fro  for  three  days,  two  species  of  ranid 
frogs,  5  of  lizards,  and  11  of  snakes  were  secured.  The  latter  were 
largely  of  fossorial  types,  among  them  a  burrowing  viper  (Atractaspis 
bibronii)  which  conformed  to  the  description  of  katangae  of  the  south- 
ern Belgian  Congo!  Only  one  frog  (Rana  o.  gribinguiensis)  and  one 
snake  (Calamclaps  u.  warreni)  could  be  cited  as  largely  sylvicoline. 

Nor  did  the  forest  appear  to  have  any  significance  as  a  refuge  for 
surviving  forest  forms  of  reptiles,  the  5  species  taken  within  its  con- 
fines being  coastal  bush  or  savanna  species.  Hornbills,  both  Bycanisies 
and  Lophoceros,  were  much  in  evidence,  but  no  small  birds  were  shot 
though  several  hours  were  spent  in  looking  for  them. 

Jumping  shrews  (Petrodromus  s.  sultani)  were  not  uncommon  on  the 
outskirts,  and  galagos  of  two  species  (G.  c.  lasiotis  and  G.  s.  zanzibari- 
cas)  were  present,  the  latter  extremely  plentiful.  The  place  was  a 
refuge  also  for  a  band  of  colobus  (C.  p.  palliatus)  and  some  blue 
monkey  (Cercopithecus  m.  monoidcs)  though  there  was  no  thought  of 
protecting  the  latter  for  whose  heads  a  reward  was  offered  as  they 
attacked  the  sisal  shoots.  Bats,  a  single  red  squirrel,  and  a  bushbuck 
complete  the  list  of  mammals  seen  in  the  forest,  though  other  species 
were  collected  in  the  surrounding  plantation. 

Magrotto  Mountain,  Tonga  Province.  5°8'  S.,  38°45'  E.   Alt.  2500  feet- 
Camped  on  a  low  hill  half-a-mile  as  the  crow  flies  west  of  the  factory 
of  MagrOtto  Estate. 

Arrived  on  June  27  and  remained  until  July  20th. 
The  first  five  days  were  largely  overclouded,  raw,  damp,  and  chilly. 
Cloud-mist  like  a  dense  fog,  swept  into  the  valley  each  evening  about 
4  p.m.  remaining  until  9  or  10  the  following  morning  when  dispersed 
by  the  frequent  rainstorms  which  swept  across  the  hills.  With  the 
advent  of  the  new  moon  the  vapour  was  reduced  to  capping  the  higher 
forested  ridges ;  the  hours  of  sunshine  increased  and  for  an  entire  week 
there  were  fewer  showers.  During  the  last  week,  however,  the  weather 
remained  dull  with  frequent  rainstorms.  Average  annual  rainfall  75 
inches ! 

Magrotto  Estate,  which  formerly  occupied  about  5000  acres  of  hill- 
tops capping  the  mountain,  is  reduced  to  half  that  size  today.  The 
more  than  a  million  coffee  trees  of  German  times  have  given  place 
to  the  West  African  oil-palm,  the  only  plantation  of  this  palm  in  all 


212  bulletin:  museum  of  comparative  zoology 

East  Africa.  My  camp  among  these  palms  was  almost  surrounded  by 
a  horseshoe-shaped  ridge  that  was  largely  forested,  a  gap  towards  the 
south  admitting  the  clouds  of  vapour  which,  during  the  southwest 
monsoon,  are  an  almost  regular  evening  phenomenon.  Just  below  the 
western  foot  of  the  camp  flows  a  river  which  rises  nearby.  To  the 
north  is  a  swampy  bottom,  part  of  whose  smothering  mat  of  vegetation 
we  cleared  in  our  search  for  frogs. 

Prior  to  1900,  or  thereabouts,  the  whole  mountain  was  heavily 
forested  until  much  was  cleared  for  coffee  planting.  During  the  Great 
War  natives  moved  in  on  the  crown  land  and  destroyed  almost  every 
tree.  The  hillside  directly  opposite  my  camp  had  been  cleared  and 
rows  of  Grevillea  planted  as  shade  trees  for  the  coffee.  When  the  Estate 
was  abandoned  during  the  war,  native  trees,  protected  by  the  Grevillea, 
sprang  up  and  are  now  between  60  and  70  feet  in  height.  I  mention 
this  as  representing  the  most  amazing  come-back  of  forest  which  I 
have  seen  anywhere  in  East  Africa.  The  forest-floor  conditions  ap- 
peared indistinguishable  to  me  from  those  in  adjacent  virgin  forest. 

Despite  this,  however,  animal  life,  which  in  species  closely  resembles 
the  fauna  of  Amani  at  3000  feet  in  the  Usambara  Mountains,  and  only 
20  miles  distant  across  the  plain,  was  decidedly  scarce;  birds  were  con- 
spicuously absent.  In  part,  of  course,  this  may  be  seasonal,  for  the 
three  weeks  spent  at  Amani  in  1926  were  during  the  November  rains 
when  amphibia  were  breeding,  hence  their  predators  more  in  evidence. 
The  three  weeks  spent  on  Magrotto  were  immediately  following  the 
coldest  months  in  the  year  when  the  temperature  falls  57°  F.  (14°  C.) 
and  a  proportion  of  poikilothermous  vertebrates  may  be  assumed  to 
be  quiescent. 

Another  factor,  though  only  affecting  the  snake  census,  might  be 
found  in  the  composition  of  the  plantation  personnel.  At  Amani  the 
'hands'  were  largelv  of  the  Nvamwezi  tribe,  who  are  notoriouslv  less 
fearful  of  these  reptiles.  At  Magrotto  the  labour  was  largely  drawn 
from  local  people,  who,  though  helpful,  and  willing  to  bring  in  such 
snakes  as  they  came  across,  displayed  no  great  enthusiasm. 

At  Magrotto  21  species  of  amphibia  were  collected  of  which  only  3 
had  not  been  taken  at  Amani,  1  of  these  was  the  forest-edge  form 
(Rana  m.  venusta)  of  the  widely  distributed  savanna  species;  signifi- 
cantly enough  both  the  others  were  recent  invaders  from  the  coastal 
plain,  i.e.  Rana  o.  oxyrhynchus  (instead  of  the  forest-edge  R.  o.  gribin- 
guiensis  taken  at  Amani)  and  Arthroleptis  s.  stenodactylus  (instead  of 
the  sylvicoline  A.  s.  lonnbergi  taken  at  Amani).  Three  remaining 
species  taken  at  Amani  though  not  encountered  on  Magrotto,  were 


loveridge:  African  itinerary  and  comments  213 

Xenopus  and  Hyperolius  forms  typical  of  the  coastal  plain  and  which 
almost  certainly  will  be  found  on  Magrotto.  A  good  series  of  topotypes 
of  H.  substriatus  (= puncticulatus)  were  preserved. 

When  we  examine  the  composition  of  Magrotto's  reptile  fauna,  of 
which  20  kinds  were  collected,  we  encounter  a  similar  situation.  In- 
stead of  Typhlops  p.  gierrai,  whose  presence  one  might  reasonably  have 
expected,  we  encountered  only  the  typical  form;  as  for  Neusterophis 
olivaceus,  the  montane  uluguruensis  occurred  in  almost  equal  propor- 
tions to  the  typical  savanna  race.  To  sum  up,  for  both  snakes  and 
lizards,  precisely  50%  were  sylvicoline,  the  rest  savanna,  reflecting 
the  fact  that,  though  we  spent  more  time  hunting  in  the  forest,  reptile 
life  was  more  conspicuous  in  the  plantation  where  immigrant  forms 
were  alike  supplanting  the  forest  fauna  in  the  zoological  and  botanical 
realms. 

As  already  stated,  forest  birds  were  scarce  at  the  time  of  our  visit, 
only  6  or  7  being  obtained,  of  which  half  were  bulbuls.  To  these  might 
be  added  the  great  eagles  (Stephanoaetus  coronatus)  seen.  Of  the  species 
collected  1  (or  2)  were  described  from  the  nearby  Usambara  Moun- 
tains, 4  from  Kilimanjaro,  and  1  from  Mt.  Elgon.  A  total  of  approxi- 
mately 25%  of  the  species  being  sylvicoline. 

Similarly  with  the  mammals  not  more  than  25%  of  the  16  species 
collected  were  forest  forms  in  its  restricted  sense,  though  the  propor- 
tion might  be  raised  almost  to  50%  by  the  inclusion  of  certain  doubt- 
ful creatures  like  Heliosciurus  u.  undulatus  of  Kilimanjaro  which,  like 
the  galagoes  and  monkeys,  are  arboreal  and  probably  as  much  at 
home  in  the  coastal  bush  as  in  virgin  rain  forest. 

Tanga,  Tanga  Province.   5°40'  S.,  39°7'  E.    Alt.  50  feet. 

At  Tanga  Hotel  prior  to  embarkation. 

Arrived  at  noon  on  July  21  and  left  on  the  23rd. 

Fine  and  hot.   Average  annual  rainfall  59.24  inches. 

Spent  my  last  afternoon  in  Tanganyika  searching  for  Typhlops 
platyrhynchus,  of  which  Tanga  is  type  locality,  in  the  sandy  area  from 
which  the  coconut  palms  had  been  removed  in  order  to  prepare  the 
site  for  a  military  landing  field.  All  that  we  got  were  3  species  of  frogs 
and  2  of  geckos,  all  typical  of  the  coastal  plain  life  zone. 

KENYA  COLONY 

Likoni,  Set/die  Province.   4°5'  S.,  39°39'  E.   Alt.  50  feet. 
On  board  R.  M.  S.  Dunbar  Castle. 
Docked  alongside  Kilindini  Wharf  July  24  and  sailed  on  the  26th. 


214  bulletin:  museum  of  comparative  zoology 

Showers  and  sunshine.  Rainfall  very  variable,  but  the  annual 
average  about  51  inches. 

Likoni  is  a  ferry  landing  on  the  mainland  opposite  Kilindini,  Mom- 
basa Island,  the  second  locality  in  bold  face  type  appearing  on  many 
labels.  It  is  a  region  of  old  coconut  plantations  and  typical  coastal 
vegetation.  I  spent  two  mornings  there  in  search  of  additional  ma- 
terial of  a  new  gecko  (Hemidactylus  t.  barbouri)  which  I  had  taken 
previously  at  nearby  Changamwe,  and  which  I  purposed  describing. 
In  addition  to  this  and  other  lizards  we  captured  the  first  three 
examples  of  a  skink  (Riopa  pembanum)  which  I  had  ever  collected,  the 
species  being  unknown  from  the  continent  until  relatively  recently. 
A  frog,  two  pigmy  mice  (Leggada  b.  vicina)  of  a  race  described  from 
Takaungu  a  few  miles  to  the  north,  and  a  good  haul  of  invertebrates 
comprised  the  final  spoils  of  a  trip  which  had  lasted  nine  months. 


PLATES 


PLATE  1 


Loveridge — African  Itinerary 


PLATE  1 

Map  showing  Principal  Collecting  Localities 

1938 

Landing  at  Mombasa  (25.x),  except  for  a  stopover  at  Naivasha  and  Kinan- 
gop  (26-31. x),  Loveridge  proceeded  by  rail  direct  to  Jinja  (1-5. xi).  Thence  to 
Mabira  Forest  (5-21.xi),  Budongo  Forest  (22.xi-7.xii),  Kibale  Forest  (8-19.xii), 
Bundibugyo  near  Bwamba  Forest  (19-26. xii),  Bugoye,  foot  of  Ruwenzori 
Mountains  (26-28.xii)  and  Mubuku  Valley  at  7000  ft.  (29.xii-). 

1939 

On  leaving  Mubuku  (-9.i)  Loveridge  descended  down  the  vallev  to  Mihunga, 
circa  6000  ft.  (9-21.i),  then  back  to  Bugoye  (21-24.i),  Nyakabande  (25-30.i), 
Mushongero  (30.i-4.xi),  returned  to  Nyakabande  (4-8.U);  Kisenyi  (8-13. ii), 
Goma  (13-14.ii),  Mamvu  on  Idjwi  Island  (14-16. ii),  Upper  Mulinga  on  Idjwi 
(16.ii-6.iii),  Uvira  (7-8.iii),  TJjiji  (9-16. iii),  Dar  es  Salaam  (18-19.iii),  Mikin- 
dani  (22-24.iii),  Mbanja  (25.iv-6.v),  Lake  Rutamba  (6-8.v),  Nchingidi 
(9-21.v),  Lindi  (22.v-4.vi),  Siga  Caves  (7-17.vi),  Amboni  Estate  (17-27.vi), 
Magrotto  Mountain  (27.vi-21.vii),  Tanga  (21-23.vii),  Kilindini  (24-26.vii). 


BULL.    MUS.    COMP.    ZOOL. 


Loveridge.    African  Itinerary  and  Comments.  Plate  1 


UGANDA.- 


'■•'BuJorrao 

h>       Forest       i  ^t 
rJ*/CibaU  F°rzZ~Apt& 

Muill/iu    Valley   Bl   Sf  JMJ\ 

?'  -Cb*  Mabira  '■ 

*■*«'  MutaXijS—-^ 
Mil     jCpn>V>°»S"° 
Nyahabaniei  -f(abale 

Lake   KlvJikk.%- 

Costermansn- 

Uvira 


BELGIAN 
CONGO 


i 


K  EN  YA 
COLONY 


TAN  G  A 
TE  R  R 


«0C 

"aS"~vtto  /t,f''WW«tJo MOMBASA 
Ambon!  EstaleY^f^t 

Stga  Cai/es'T     Q 


-v  ( 9 


x/ 


* 


A 


* 


V 


ZIBAR 
ES    SALAAM 


'Mindani 


Scale  in  Miles 


0 

u. 


100 

_l_l 


200 

1 


PLATE  2 


Loveridge— African  Itinerary 


PLATE  2 

Fig.  1,     Transport  by  water — Canoes  on  Lake  Mutanda 

These  dugouts,  waiting  to  take  us  from  Mushongero  to  the  south  end  of  the 
Lake,  speak  volumes  for  the  patient  toilers  who,  with  adze  or  other  simple 
tool,  gouged  out  the  hard  timber  from  the  fallen  tree.  Moreover,  as  no  trees 
of  sufficient  height  to  provide  sixteen  or  eighteen-foot  canoes  are  to  be  found 
within  many  miles  of  the  Lake,  these  incredibly  heavy  craft  had  to  be  pushed 
uphill  and  down  dale  on  rollers  by  human  muscle  alone. 

Fig.  2.     Transport  by  Land — Our  Lorry  at  Mubango 

African  travel  today  involves  increasing  use  of  modern  methods  of  transport, 
resort  to  head-porterage  being  reserved  for  roadless  regions  or  where  the  ascent 
of  a  mountain  by  native  track  is  called  for.  In  modernizing  Africa  motor- 
driven  vehicles  will  soon  be  claiming  precedence  over  predators  and  snakes 
as  the  major  menace  to  life  and  limb.  The  photograph  depicts  our  native-driven 
Mercedes-Bentz,  which  at  the  time  was  carrying  my  six  boys  atop  a  load  con- 
sisting of  a  ton  of  camp  and  collecting  equipment,  irresolutely  resting  on 
marshy  ground  where  faulty  driving  had  landed  her  at  the  very  start  of  our 
'safari!' 


BULL.    MUS     COMP.    ZOOL. 


Loveridge.     African  Itinerary  and  Comments.  Plate  2 


PLATE  3 


Loveridge — African  Itinerary 


PLATE  3 

Fig.  1.     Prospecting  For  a  Camp  Site — Forest-edge,  Budongo 

Budongo  Forest,  which  covers  about  180  square  miles,  gathers  about  its 
fringes  a  dense  and  often  impenetrable  undergrowth  of  bush.  Within,  however, 
where  the  unbroken  forest  canopy,  two-hundred  feet  overhead,  prevents 
shrubs  springing  up  among  the  mighty  buttress  roots,  one  can  wander  at  will 
over  the  thick  layer  of  leaves  which  carpet  the  forest  floor. 

Fig.  2.     Tractor  and  Tramway  are  Employed  for  Logging  at  Bisu 

The  absence  of  paths  and  water  drove  us  unwillingly  to  camp  at  Bisu,  in 
close  proximity  to  the  Buchanan  Saw  Mills.  There  access  to  the  forest  was 
assured  by  a  track  cut  for  the  tramway  depicted  above.  Unfortunately  the 
noisy  tractor  and  its  train,  which  made  the  three-mile  run  through  the  forest 
every  twelve  hours,  had  driven  elephant,  buffalo,  and  other  large  mammals  to 
seek  quiet  elsewhere. 


BULL.    MUS.    COMP.    ZOOL. 


Loveridge.     African  Itinerary  and  Comments.  Plate  3 


I 


MHMETttBl^BUHB 


PLATE  4 


Loveridoe — African  Itinerary 


PLATE  4 

Fig.  1.     Native  Path  through  Mabira  Forest 

Paths  were  few  and  far  between,  yet  the  undergrowth  was  so  impenetrable 
that  they  provided  the  only  means  for  reaching  the  deeper  forest  to  sample  its 
denizens.  While  monkeys  and  squirrels  fell  victims  to  this  inquisitiveness, 
parrots  screeched  or  clambered  about  in  the  tree-tops,  where,  with  a  host  of 
smaller  birds  feeding  in  the  forest  canopy,  they  enjoyed  complete  immunity, 
being  out  of  range  of  gunshot. 

Fig.  2.     A  Semi-pigmy  of  Baamba  or  Batwa  Stock 

The  Baamba  clans  inhabiting  the  Ituri  Forest  region  northwest  of  the 
Ruwenzori  Mountains,  closely  resemble  their  Batwa  kinsmen  of  the  Congo 
forests.  Attired  only  in  a  civet  skin,  the  sturdy  subject  of  the  photograph 
presents  a  marked  contrast  to  the  tall,  cotton-clad  Muganda  on  the  forest 
path  in  Mabira. 

Fig.  3.     Family  Group  at  Bundibugyo  in  Toro 

Yet  another  popular  style  in  dress  is  exhibited  by  this  Bantu  family  relaxing 
after  their  early  morning  labours  in  the  hot  region  just  above  the  Semliki 
Valley.  Hundreds  of  natives  representing  many  tribes,  visited  our  camp  at 
Bundibugyo.  If  I  am  not  mistaken  those  shown  are  Batoro,  the  agricultural 
middle-class  members  of  this  society,  superior  to  the  hunting  Baamba  from 
whom  they  obtain  meat  in  exchange  for  cereals,  yet  themselves  formerly  sub- 
ject to  their  pastoral  overlords,  the  Hamitic  Bahima. 


BULL.    MUS.    COMP.    ZOOL 


Loveridge.     African  Itinerary  and  Comments.  Plate  4 


r-£.j 


# 


■& 


■ 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  6 


NOTES  ON  LYCAENID  BUTTERFLIES 


By  Harry  K.  Clench 


MUS.  COMP.  ZOOL 
LIBRARY 

SEP  2  2  1964 

HARVARD 
UNIVERSITY. 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED  FOR  THE  MUSEUM 

July,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2, 
3,  4  and  5  have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


SEP  2  2  1964 


HARVARD 
Bulletin  of  the  Museum  of  Comparative  Zoology     rsiTY 

AT  HARVARD   COLLEGE 

Vol.  XCIV,  No.  G 


NOTES  ON  LYCAENID  BUTTERFLIES 


Bv  Harry  K.  Clench 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED   FOR  THE  MUSEUM 

July,  1944 


JUL  |9  1944 


L  I  B  K  A  K  * 


No.  G  —  Notes  on  Lycaenid  Butterflies 
By  Harry  K.  Clench 

a.  THE  GENUS  CALLOPHRYS  IN  NORTH  AMERICA 

While  working  on  a  revision  of  the  genus  Indsalia  Scudder  sueh 
frequent  recourse  was  had  to  the  various  American  species  of  Callo- 
phrys  that  it  was  thought  advisable  to  prepare  a  brief  review  of  them. 
Unfortunately,  time  was  all  too  short  to  do  the  job  properly,  and 
genitalic  examinations  were  not  made.  These  notes,  however,  were 
gotten  together,  in  the  hope  that  they  would  facilitate  further  work 
on  the  genus.  Such  future  work  should  certainly  include  a  genitalic 
study  of  all  forms  involved. 

The  first  and  only  real  revision  of  the  American  Callophrys  was  that 
of  Barnes  and  Benjamin' ,  who  placed  the  genus  on  very  solid  footing, 
especially  as  compared  with  its  previous  state.  It  was  through  this 
paper  that  our  first  accurate  knowledge  of  the  limits  and  variabilities 
of  the  several  species  was  obtained.  There  have  remained,  however, 
several  points  that  need  clarification.  Probably  the  most  important 
of  these  is  the  interrelationship  of  the  various  Californian  forms.  The 
arrangement  adopted  by  Barnes  and  Benjamin  (dumetorum  in  the 
north,  with  subsp.  perple.va  in  the  south)  is  too  simple,  and  although 
it  at  first  appears  logical,  subsequent  research  has  proved  it  false. 
There  are  at  least  three  forms,  and  very  likely  several  more,  in  Cali- 
fornia that  have  been  going  under  the  names  dumetorum  and  perplexa. 
What  has  been  commonly  passing  as  true  dumetorum  is  something 
else,  while  true  dumetorum  itself  appears  to  be  rather  poorly  known. 
These  will  be  discussed  in  greater  detail  under  the  species  concerned. 
Unfortunately,  the  problem  in  California  has  been  only  partly  solved, 
the  material  at  hand  indicating  much  further  work  to  be  done,  but  of 
itself  insufficient  to  do  it. 

In  addition  to  this,  there  has  been  a  new  species  (comstocJci)  pro- 
posed since  the  work  of  Barnes  and  Benjamin,  and  in  the  present 
paper  a  second  novelty,  a  new  subspecies  of  affinis,  is  added. 

Finally,  there  are  a  number  of  new  and  interesting  records,  exten- 
sions of  ranges,  particularly  of  affinis  and  sheridanii. 

The  genus  in  North  America  appears  to  be  confined  exclusively  to 
the  Rocky  Mountains  and  the  area  westward  to  the  Pacific.  It  ex- 
tends from  northern  Mexico  in  the  south  to  Alberta  and  British 

U923,  Contrib.  Nat.  Hist.  Lep.  North  America  5,  p.  61-67. 


218  bulletin:  museum  of  comparative  zoology 

Columbia1  in  the  north.  The  species  appear  to  be  generally  mountain- 
loving,  although  they  are  frequently  (perplexa,  for  example)  taken  at 
low  elevations. 

The  paucity  of  species  in  Asia  and  Europe  and  their  close  inter- 
similarity,  as  compared  with  the  relatively  large  (6)  number  of  species 
and  their  diversity  here  in  America,  would  suggest  that  the  Palae- 
arctic  species  were  derived  from  American  stock.  The  origin  of  this 
American  stock,  however,  is  quite  another  question,  and  one  foolish 
to  speculate  upon,  in  view  of  our  present  ignorance  of  much  of  the 
pertinent  details. 

The  author  wishes  to  thank  the  several  individuals  and  institutions 
who  have  greatly  assisted  in  this  review.  The  American  Museum  of 
Natural  History  (AMNH)  very  kindly  loaned  some  material  for 
study.  The  Carnegie  Museum  and  the  U.  S.  National  Museum 
checked  the  types  of  species  in  their  collections.  Mr.  Robert  G.  Wind, 
of  Berkeley,  California,  was  very  kind  in  sending  me  for  study  a  large 
amount  of  exceedingly  interesting  material,  including  a  number  of  the 
complex  viridis-\\ke  specimens  from  the  Californian  Sierras,  and  other 
northern  Californian  localities.  Mr.  Charles  L.  Remington  also  lent 
some  very  interesting  material  which  was  of  the  utmost  help  in  pre- 
paring the  paper.  Mr.  L.  P.  Grey  gave  the  author  all  of  his  Callophrys, 
which  included  several  very  interesting  things.  The  collection  of  the 
Museum  of  Comparative  Zoology  (M.C.Z.)  has,  as  in  times  past, 
always  been  available  to  me  through  the  kindness  of  Mr.  Banks.  This 
collection  contains  much  material  of  interest  and  assistance.  My  own 
collection  has  been  drawn  on  wherever  possible,  being  designated 
(H.K.C.). 

Genus  Callophrys  Billberg 

1820,  Enumeratio  Insectorum,  p.  80.    Genotype,   Papilio  rubi  Linn.     (Palae- 
arctic). 

Key  to  species 

1.  Fore  wing  underside  with  green  covering  wing  down  to  C112  or  2A 2 

-    This  green  restricted  to  a  more  or  less  broad  costal  and  outer  marginal 
border,  leaving  a  large  internal  area  of  gray  or  fulvous 6 

'The  two  papers  on  the  faunae  of  these  provinees  (K.  Bowman,  1919,  Annotated  Cheek  List 
of  the  Macrolepidoptera  of  Alberta,  Alberta  Nat.  Hist.  Soc.,  Red  Deer,  Alta.,  16  pp.;  E.  H. 
Blackmore,  1927,  Check-list  of  the  Macrolepidoptera  of  British  Columbia,  C.  F.  Banfield, 
Victoria,  B.  C  47  pp.)  each  list  Callophrys  dumelorum.  Just,  what  they  meant  is  not  known. 
Indeed,  it  coidd  have  been  almost  any  species  in  the  genus,  but  most  logically  affinis  or  shcri- 
danii.  Beferences  to  these  papers,  as  well  as  to  a  number  of  others,  were  omitted  due  to  this 
and  similar  lack  of  precise  information,  so  common  in  this  genus. 


clench:  notes  on  lycaenid  butterflies  219 

2.  Hind  wing  below  immaculate,  or  at  most  with  a  faint  indication  of  a  white 

line — usually  a  discal  row  of  obscure  white  dashes 3 

-  This  wing  with  either  a  prominent  white  line,  or  with  a  series  of  distinct, 

pure  white  dashes  or  dots 4 

3.  Fulvous  above  in   both  sexes  widespread,   dominant;   underside   apple- 

green  a.  ajfmis 

-  Fulvous  above  reduced,  frequently  (usually?)  absent  in  the  male;  green 

below  purer,  inclining  even  to  bluishness affinis  washingtonia 

4.  Underside  of  hind  wing  with  a  solid  line  of  white 5 

-  This  surface  with  a  row  of  prominent  white  spots viridis 

5.  Line  on  underside  of  hind  wing  heavy,  inwardly  edged  with  a  pronounced 

band  of  black s.  sheridanii 

-  This  line  thin;  the  black  faint  or  absent sheridanii  neoperplexa 

6.  Underside  of  hind  wing  nearly  immaculate,  or  with  a  row  of  white  spots 

of  varying  number 7 

-  This  area  with  a  crooked  white  line 8 

7.  Outer  margin  of  fore  wing  evenly  convex apama  homoperplexa 

-  This  margin   convexly   angled  at   M2  or  thereabouts,   and  straight,    or 

slightly  concave  thence  to  the  inner  angle 9 

8.  Line  on  underside  of  hind  wing  basally  edged  with  black,  then  fulvous 

a.  apama 

-  This  line  with  no  fulvous comstocki 

9.  Costa  of  fore  wing  below  edged  with  fulvous dumetorum  perplexa 

-  Costa  of  this  surface  not  edged  with  fulvous  (or  at  most  only  very  slightly) 

d.  dumetorum 


Callophrys  dumetorum  Boisduval 

This  species  ranges  from  northern  Mexico  (Baja  California)  north 
beyond  San  Francisco.  The  northern  limits  of  its  range  are  still  un- 
defined. There  are  apparently  two  subspecies  of  this,  one  very  widely 
known  (perplexa),  inhabiting  the  lower  part  of  its  range,  and  the  other 
(typical  dumetorum)  much  less  perfectly  known,  inhabiting  the  upper. 
It  is  possible  that  the  latter  as  considered  herein  consists  in  reality  of 
several  subspecies. 

The  species  shows  several  characters  that  place  it  close  to  apama. 
The  principal  one  of  these  is  the  large  fulvous  or  gray  area  on  the  fore 
wing  below.  The  chief  constant  difference  between  the  two  (apama 
homoperplexa  and  dumetorum  perplexa  approach  each  other  so  closely 
that  aside  from  this  basic  character  they  are  almost  inseparable)  lies 
in  the  shape  of  the  outer  margin.  In  dumetorum  it  is  convexly  angled 
in  the  vicinity  of  M2,  and  straight  or  slightly  concave  below  it  to  the 
inner  margin,  while  in  apama  the  whole  margin  is  evenly  convex. 


220  bulletin:  museum  of  comparative  zoology 

Callophrys  dumetorum  dumetorum  Boisduval 

Thecla  dumetorum  Boisduval,  1852,  Ann.  Soc.  Ent.  France  (2)  10,  p.  291; 
Oberthur,  1913,  Et.  Lep.  Comp.  9,  p.  40,  pi.  236,  fig.  1926;  Draudt,  1919, 
in  Seitz,  Macrolep.  World,  6,  p.  763,  pi.  154b;  id.,  1924,  p.  1043.  (all 
partim) 

Thecla  dumetorum:  auct.  (partim) 

Callophrys  dumetorum:  auct.  (partim) 

The  type,  in  the  United  States  National  Museum,  came  from  Cali- 
fornia. No  definite  locality  was  designated.  Due  to  a  lack  of  material 
agreeing  satisfactorily  with  the  description  and  with  Oberthur's 
figure  of  the  type  (loc.  cit.)  no  type  locality  has  been  fixed. 

Localities.  California:  Phelan;  Snow  Creek;  Dana  Point  (all 
H.K.C.);  Pasadena  (M.C.Z.);  Calistoga;  Petrified  Forest  (both  in  the 
collection  of  R.  G.  Wind).  These  are  all  doubtfully  typical  dumetorum. 

Male  above  uniform  brownish  gray.  Fringe  white,  basally  gray. 
Female  similar  but  more  brownish,  and  with  a  discal  fulvous  suffusion. 

Male  below  with  fore  wing  gray  or  fulvous,  inner  margin  somewhat 
paler.  Costa  and  outer  margin  as  far  down  as  Cui  green.  Hind  wing 
uniform  green  with  two  or  three  white  spots,  the  principal  ones  being 
on  the  costa  and  in  the  CU1-CU2  interspace.  Fringe  gray,  slightly 
paler  outwardly  and  at  the  vein-ends  on  the  hind  wing.  Female 
similar,  but  with  the  gray  area  on  the  fore  wing  more  fulvous.  Both 
sexes  frequently  have  a  more  or  less  strongly  developed  discal  line  on 
the  fore  wing,  usually  merely  an  intensification  of  the  gray  or  fulvous. 

Length  of  fore  wing.    Male,  12.5-13  mm.;  female,  13  mm. 

The  above  rather  brief  description  is  based  on  three  males  and  a 
female  (two  males  from  Phelan,  one  from  Snow  Creek,  and  a  female 
from  Dana  Point)  which  may  or  may  not  be  typical  dumetorum. 
Several  points  in  Boisduval's  description  differ  from  those  found  in 
these  specimens  (namely,  the  fulvous  (of  Boisduval)  instead  of  gray 
color  of  the  large  area  on  the  fore  wing  below;  the  presence  of  a  number 
of  white  spots  on  the  hind  wing  below,  lacking  in  the  above-described 
specimens).  Typical  dumetorum  was  very  likely  described  from  much 
further  north  than  these  four  specimens,  although  just  where  is  still 
unknown.  The  Calistoga  specimens,  while  apparently  approaching 
Boisduval's  description  a  little  more  closely  than  these,  still  differ 
enough  to  be  closer  to  the  more  southern  specimens. 

These  specimens,  of  what  is  here  taken  to  be  typical  dumetorum, 
differ  from  perplexa  in  the  absence  of  fulvous  edging  on  the  costa  of  the 
fore  wing  below,  and  (in  the  male)  by  the  gray,  instead  of  fulvous, 
area  on  this  surface.  There  are,  however,  hints  of  fulvous  on  this  area, 


clench:  notes  on  lycaenid  butterflies  221 

but  this  may  possibly  be  explained  by  the  fact  that  the  specimens 
came  from  a  region  not  too  remote  from  the  home  of  true  perplexa. 

Two  of  the  three  ( lalistoga  males  examined  have  a  definite  fulvous 
shading  on  the  hind  wing  above,  and  one  of  them  a  patch  on  the  fore 
wing  as  well.  More  specimens  are  needed  to  tell  whether  or  not  this  is 
a  local  race  of  dumetorum.  This  character  is  quite  unusual  for  dume- 
torum  and  appears  not  to  have  been  noticed  previously. 

Callophrys  dcmetorum  perplexa  Barnes  and  Benjamin 

Thecla  affinis:  auct.  (partim) 

Thecla  dumetorum:  auct.  (partim) 

Callophrys  dumetorum:  McDunnough,  1914,  Ent.  Rec.  and  Journ.    Variation, 

26,  p.  196.    (partim) 
Callophrys  dumetorum  race  perplexa  Barnes  and  Benjamin,   1923,   Contrib. 

Nat.  Hist.  Lep.  North  America,  5,  p.  65;  Draudt,  1924,  in  Seitz,  Macrolep. 

World,  5,   p.    1043.    Comstock,   1927,   Butterflies  of  California,   p.    168, 

pi.  50,  figs.  17,  18,  19  (partim);  Hoffman,  1940,  Anales  Inst.  Biol.  Mex.,  11, 

p.  708  (no.  623). 

Holotype,  allotype  and  a  number  of  paratypes  in  the  United  States 
National  Museum.   Described  from  San  Diego,  California. 

Other  localities.  California :  Pine  Valley ;  Corona ;  Balboa ;  El  Modena 
(all  in  coll.  H.K.C.).  Mexico:  (northern  localities,  adjacent  to  Cali- 
fornia). 

Differs  from  typical  dumetorum  (as  considered  in  this  paper)  in  the 
fulvous  edging  on  the  costa  of  the  fore  wing  below,  and  in  the  in- 
creased fulvous  in  the  discal  region  of  this  same  surface. 

Apparently  restricted  to  southern  California  and  adjacent  Mexico. 

Length  of  fore  wing.   Male,  11-13.5  mm.;  female,  11.5  mm. 

Callophrys  comstocki  Henne 
Callophrys  comstocki  Henne,  1940,  Bull.  So.  Cal.  Acad.  Sci.,  39,  p.  71. 

Holotype  and  allotype  from  the  Providence  Mts.,  San  Bernardino 
County,  California,  in  the  collection  of  the  Los  Angeles  County 
Museum.  Paratypes  from  the  same  locality  in  the  United  States 
National  Museum,  Canadian  National  Collection.  Also  (?)  in  the 
collections  of  C.  H.  Ingham  and  C.  Henne. 

Topotypes  have  been  examined  from  the  collections  of  A.  C. 
Frederick,  D.  B.  Stallings  and  the  author. 

Above,  both  sexes  slate  gray.  Male  with  a  small  oval  scent-pad. 
Fringe  gray,  paler  outwardly.    Hind  wing  but  slightly  scalloped. 


222  bulletin:  museum  of  comparative  zoology 

Below,  both  sexes  flat  grayish  green.  Fore  wing  with  inner  marginal 
area,  centrally  as  far  costad  as  M3,  gray.  This  wing  crossed  by  an 
obsolescent  post-discal  line  of  white,  basally  gray-black,  dashes. 
Hind  wing  with  a  similar,  but  more  distinct,  line,  edged  basally  with 
black.  This  line  outwardly  displaced  in  M3-C111,  and  irregular  thence 
to  the  inner  margin.  Fringe  of  both  wings  basally  gray,  outwardly 
white. 

Length  of  fore  wing.   Male,  10-11  mm. 

Closest  to  apama,  but  differing  in  several  respects,  principally  the 
gray  upper  surface  of  both  sexes,  and  the  flat  green  below,  unrelieved 
by  any  fulvous  shading.  It  is  a  very  interesting  and  isolated  species, 
possibly  a  local  modification  of  apama,  but  certainly  distinct  enough 
to  warrant  full  specific  distinction.  So  far  as  known  it  is  restricted  to 
the  Providence  Mts. 

Callophrys  apama  Edwards 

This  species  occurs  over  a  relatively  compact  range  from  Arizona 
to  New  Mexico  and  north  to  Colorado,  becoming  differentiated  in  the 
latter  regions  into  a  distinct  subspecies.  The  limits  of  range  of  this 
species  are  fairly  well-known.  It  may,  however,  be  turned  up  in 
eastern  California  (doubtful)  or  southern  Utah.  (The  Carnegie 
Museum  has  a  single  male  from  this  state;  I  have  not  seen  it.) 

From  its  near  relative,  C.  dianetorum,  apama  may  be  distinguished 
by  the  character  given  above  under  the  general  remarks  for  dume- 
torum. 

Another  close  ally  (perhaps  its  closest)  is  the  recently  described 
C.  comstocki.  This  latter  may  be  separated  at  once  by  its  complete 
lack  of  fulvous  shading  on  the  underside,  by  its  smaller  size,  and  by 
the  more  distorted,  less  prominent  white  discal  line  on  the  hind  wing 
below. 

Callophrys  apama  apama  Edwards 

Thecla  apama  Edwards,  1882,  Papilio,  2,  p.  137;  Draudt,  1919,  in  Seitz,  Macro- 

lep.   World,  5,  p.  763;  id.,  1924,  p.  1043. 
Callophrys  apama:  Barnes  and   Benjamin,    1923,   Contrib.   Nat.   Hist.   Lep. 

North  America,  5,  p.  67. 

Type  locality.  Fort  Grant  in  the  Graham  Mts.,  Arizona.  One  male 
and  two  females  labelled  "Arizona"  in  the  Edwards  collection  at  the 
Carnegie  Museum.   One  of  the  latter  made  a  lectotype  by  Holland. 

Other  localities.  Arizona:  Oak  Creek  Canyon  (6000  ft.)  (M.C.Z.); 
White  Mts.;  Catalina  Mts.  (both  in  coll.  H.K.C.);  Navajo  Mt.  (in 


clench:  notes  on  lycaenid  butterflies  223 

coll.  R,  G.  Wind);  Sierra  Aneha  Mts.  (in  coll.  P.  S.  and  C.  L.  Reming- 
ton). 

Above  in  the  male  dark  gray  with  a  brownish  tinge.  A  small, 
almost  circular  scent-pad  at  the  upper  cell-end.  Fringe  of  fore  wing 
concolorous  with  ground  (or  slightly  darker),  paler  outwardly;  of 
hind  wing  similar,  but  frequently  almost  pure  white  outwardly;  tips  of 
veins  Cuu  Cu2j  and  2A  usually  white. 

Female  similar,  but  with  fulvous  patch  on  fore  wing  and  usually 
on  hind  wing.   Fringe  outwardly  slightly  paler. 

Below,  fore  wing  largely  fulvous.  Inner  margin  gray.  A  post- 
discal,  white  irregular  line  crosses  from  costa  to  Cu2,  composed  of 
white  dashes,  basally  edged  with  black.  Base  of  wing  and  apical  area 
outside  of  post-discal  line  (and  extending  down  as  far  as  Cux)  green. 
Hind  wing  green.  A  post-discal  line,  similar  in  construction  to  that  of 
the  fore  wing,  but  here  more  tortuous,  crosses  from  costa  to  inner 
margin,  outwardly  displaced  often  in  the  form  of  a  crude  "\Y"  in  the 
M3-Q12  region.  Basal  to  this  line  runs  a  closely  adherent  band  of 
fulvous.  Occasionally  there  is  a  definite  suggestion  of  a  Cui-Cu2  sub- 
marginal  spot.  Fringe  of  fore  wing  basally  dull  fulvous,  outwardly 
gray;  of  hind  wing  similar,  but  paler  outwardly  and  tipped  at  Ciii,  Cu2 
and  2 A  with  white. 

Length  of  fore  wing.   Male,  10.5-13  mm.;  female,  10.5-11  mm. 

The  typical  subspecies  is  distinguished  from  its  Colorado  repre- 
sentative by  the  clarity  and  completeness  of  the  markings  below, 
which  are  reduced  frequently  to  the  point  of  complete  obscurity  in 
homoperplexa. 

Callophrys  apama  homoperplexa  Barnes  and  Benjamin 

Thccla  dumetorum  auct.  (partim) 

Callophrys  apama  race  homoperplexa  Barnes  and  Benjamin,   1923,   Contrib. 

Nat.  Hist.  Lep.  North  America,  5,  p.  68;  Draudt,  1924,  in  Seitz,  Macro- 

lep.   World,  5,  p.  1043. 

Holotype,  allotype,  and  several  paratypes  in  the  United  States 
National  Museum.  Holotype  and  allotype  from  Denver,  Colorado. 
Paratypes  from  Golden,  Boulder,  and  Denver,  Colorado. 

Other  localities.  Colorado:  Durango;  Husted  and  Starr  Ranch 
(El  Paso  County)  (all  M.C.Z.);  Boulder  Canyon;  Eldora  (both  in 
coll.  P.  S.  and  C.  L.  Remington).  New  Mexico:  nr.  Hot  Springs,  Las 
Vegas  (7000  ft.);  Rincon  (both  in  M.C.Z.).  Specimens  from  Golden 
have  also  been  examined. 


224  bulletin:  museum  of  comparative  zoology 

On  the  upperside  it  differs  apparently  only  in  the  more  fulvous 
shading  of  the  female,  and  a  greater  tint  of  fulvous  in  the  male. 

On  the  underside  the  differences  are  much  more  pronounced. 
Homopcrple.va  lacks,  in  varying  stages  of  completeness,  the  white 
discal  lines  on  each  wing  which  form  such  a  conspicuous  part  of  typical 
apama.  The  green  on  the  fore  wing  appears  to  be  more  greatly  ex- 
tended. 

Length  of  fore  wing.   As  in  the  typical  subspecies. 

This  subspecies  is  very  interesting,  and  would  seem  to  point  to- 
wards a  connection  with  affinis.  This  at  first  implausible  suggestion 
becomes  more  likely  when  one  notices  that  males  of  this  subspecies  are 
frequently  dull  fulvous,  and  that  occasionally  specimens  are  found  in 
which  the  green  of  the  underside  of  the  fore  wing  covers  an  abnormally 
large  part  of  the  wing,  or  shows  itself  faintly  as  a  discal  green  suffusion. 
Another  point  in  favor  of  this  alliance  is  the  tendency  of  affinis  to 
produce  specimens  with  hints  (sometimes  startlingly  complete)  of  a 
discal  white  line  on  the  hind  wing  below.  Yet  another  point  is  that  the 
ranges  of  the  two  do  not  apparently  overlap,  or  if  so,  they  only  occur 
sympatrically  in  a  very  restricted  area.  More  material  is  needed  from 
the  zone  of  transition  (northern  Colorado  and  adjacent  regions)  to 
prove  or  disprove  this. 

Prior  to  the  work  of  Barnes  and  Benjamin,  this  subspecies  was  com- 
pletely confused  with  dumetorum  perple.va  (dumetorum  auct.)  of 
southern  California,  and  very  understandably  so.  The  two  are  strik- 
ingly similar,  and  frequently  almost  the  only  real  point  of  distinction 
(superficial)  is  the  difference  in  the  fore  wing  outline.  In  general,  how- 
ever, males  of  homoperplexa  have  a  more  ruddy  upper  surface  than  do 
males  of  perple.va.  The  green  below  of  homoperplexa  tends  to  be  some- 
what more  brassy  than  that  of  perple.va. 

Extreme  homoperplexa  lacks  any  suggestion  of  the  discal  line  on 
either  wing  below.  Throughout  Colorado,  however,  and  particularly 
in  the  southern  part  of  the  state  specimens  are  frequently  found  that 
tend  quite  definitely  toward  the  typical  subspecies.  The  two  New 
Mexico  examples  (localities  given  above)  are  in  this  category  as  well. 
Apparently  apama  is  restricted  to  Arizona. 

Callophrys  affinis  Edwards 

The  exact  range  of  this  species  is  not  yet  known.  It  extends,  from 
the  material  and  information  at  present  available,  from  Utah  north  to 
Wyoming  and  Washington.    It  may  possibly  occur  in  Colorado  also. 


clench:  notes  on  lycaenid  butterflies  225 

The  Washington  specimens  appear  sufficiently  distinct  to  warrant 
separate  racial  consideration.  Wyoming  specimens  appear  to  be 
transitional  in  some  respects,  but  are  referred  at  present  to  the  typical 
subspecies. 

This  species  is  distinguished  by  its  large,  usually  entirely  unmarked 
green  under  surface,  and  (with  the  exception  of  the  Washington  race) 
large  amount  of  fulvous  above  in  the  male.  Both  dumetorum  (Calis- 
toga)  and  apama  homoperplexa  have  this  suffusion  in  the  males,  but 
not  to  the  extent  of  typical  affinis. 

Callophrys  affinis  affinis  Edwards 

Theda  affinis  Edwards,  1862,  Proc.  Acad.  Nat.  Sri.  Philadelphia,    p.  223; 

Draudt,  1919,  in  Seitz,  Macrolep.  World,  5,  p.  763;  id.,  1924,  p.  1043. 
Callophrys  affinis:  Barnes  and  Benjamin,  1923,  Contrib.  Nat.  Hist.  Lep.  North 

America,  5,  p.  66. 

The  types,  one  male  and  one  female,  from  Utah,  are  in  the  Carnegie 
Museum.  Silver  Lake,  Utah,  is  here  selected  as  the  type  locality, 
based  upon  two  males  and  two  females  in  the  M.C.Z.  Barnes  and 
Benjamin  (loc.  cit.)  referred  to  some  specimens  from  Silver  Lake  as 
topotypes,  but  did  not  elaborate. 

Other  localities.  Wyoming:  Teton  Mts.  (in  coll.  R.  G.  Wind). 
Edwards'  description  of  this  species  is  very  good,  and  is  quoted  here: 
"Both  sexes  glossy  red  brown,  brightest  in  female;  the  male  has  a 
smooth  oval  spot  on  disc  of  primaries;  costa  of  primaries  and  base  of 
both  wings  blackish  brown;  whole  hind  margin  edged  with  same 
color;  fringe  white. 

"Under  side  uniform  apple  green,  except  on  inner  margin  of  pri- 
maries, where  it  is  pale,  brownish  grey;  both  wings  immaculate;  costal 
edge  of  primaries  grey;  hind  margin  of  secondaries  without  crena- 
tions." 

Length  of  fore  wing:  Male,  12-13  mm.;  female,  12.5-13.5  mm. 

The  typical  form  is  large  and  bright.  It  differs  from  washingtonia 
in  the  shade  of  apple  green  below,  and  in  the  large  extent  of  fulvous 
on  the  male  above. 

t 

Callophrys  affinis  washingtonia,  new  subspecies 

Upperside: 

Male.  Both  wings  dark  gray,  slightly  brownish,  and  occasionally 
suffused  with  dull  fulvous  in  the  disk.  Fringe  white,  basally  dark 
gray. 


226  bulletin:  museum  of  comparative  zoology 

Female.  Similar  to  the  male,  but  with  the  fulvous  more  extended. 
Underside: 

Male.  Both  wings  green,  slightly  bluish,  and  very  faintly  brassy. 
Inner  margin  of  fore  wing  to  2A,  and  frequently  to  C112,  gray.  Hind 
wing  occasionally  with  the  merest  suggestion  of  a  spot  in  the  CU1-CU2 
interspace.1  Fringe  white,  slightly  darker  at  the  vein-ends  on  the 
hind  wing. 

Female.  Similar  to  the  male,  but  with  the  addition  of  a  very  thin, 
extreme  marginal  line  of  white,  running  from  anal  angle  to  Mi.  This 
is  very  likely  an  individual  variant,  and  will  probably  not  be  found  to 
hold  true  for  other  females.  At  the  anal  angle  is  a  very  small  white 
patch,  the  origin  of  the  white  line,  and  probably  also  merely  an 
individual  variation. 

Length  of  fore  wing.   Male,  11-12.5  mm.;  female,  12  mm. 

Holotype.  Male,  Alta  Lake,  Washington,  April  25,  1935  (J.  C. 
Hopfinger)  ex  coll.  P.  S.  and  C.  L.  Remington. 

Allotype.   Female,  same  data  as  holotype. 

Paratypes.  Two  males,  Brewster,  Washington,  May  9,  1939  (J.  C. 
Hopfinger?),  ex  coll.  L.  P.  Grey. 

Holotype  and  allotype,  no.  26259  in  the  M.C.Z.  Paratypes  in  the 
author's  collection. 

Remarks.  Differs  from  typical  affinis  in  the  much  more  reduced 
fulvous  above  in  both  sexes,  and  in  the  more  bluish  tone  of  the  green 
below. 

A  series  of  over  15  examples  from  the  Teton  Mts.,  Wyoming,  seems 
about  intermediate  between  washingtonia  and  true  affinis.  There  is, 
however,  little  if  any  variation  in  the  amount  of  fulvous  on  the  fore 
wing  above  in  the  male,  while  this  character  in  washingtonia  varies 
from  a  limited  amount  (smaller  than  the  smallest  .of  the  Teton  Mts. 
specimens)  to  none  at  all.  In  several  of  the  Wyoming  specimens 
there  is  an  indication  of  a  tendency  to  produce  a  row  of  white  dashes 
on  the  hind  wing  below;  in  one  specimen  quite  strongly. 

Callophrys  viridis  Edwards 

Thecla  viridis  Edwards,  1862,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  p.  223. 

Thecla  dumetorum:  auct.  {partim) 

Callophrys  dumetorum:  Barnes  and  Benjamin,  1923,  Contrib.  Nat.  Hist.  Lep. 
North  America,  5,  p.  64  {partim);  Comstock,  1927,  Butterflies  of  Cali- 
fornia, p.  168,  pi.  50,  figs.  16,  20,  21,  25  (some  of  these  are  apparently 
dumetorum)  {partim). 

1  The  allotype  has  an  indication  of  dashes  in  other  neighboring  interspaces,  as  well  as  a  row 
of  very  faint  dashes  on  the  fore  wing.  This  latter  can  be  seen  faintly  in  certain  lights  in  one 
of  the  paratypes. 


clench:  notes  on  lycaenid  butterflies  227 

The  type  locality  is  here  selected  as  San  Francisco,  California, 
based  on  a  female  in  the  collection  of  P.  S.  and  C.  L.  Remington 
(May  8,  1934,  Wm.  Hovanitz  collector)  that  agrees  excellently  with 
the  original  description.  In  the  absence  of  any  type  at  the  Carnegie 
Museum  this  specimen  is  here  made  the  neoholotype. 

Other  localities.    "California"  (M.C.Z.). 

The  male  is  above  uniform  gray.  Fringe  white,  gray  basally  and  at 
the  vein-ends  of  the  hind  wing.  Female  similar,  but  may  be  largely 
suffused  with  fulvous,  leaving  a  dark  base,  costa  and  outer  margin  on 
the  fore  wing,  and  a  dark  base  only  on  the  hind  wing. 

Below  in  both  sexes  both  wings  are  uniform  pea-green.  Fore  wing 
with  inner  marginal  area  to  vein  Cu2  gray,  save  on  outer  margin, 
where  the  green  extends  down  to  2A.  Costa  edged  with  fulvous.  In 
the  disk  is  a  faint  row  of  dull  white  dashes,  three  in  number  in  the 
specimen  examined,  in  interspaces  M2-M3-Cu1-Cu2.  On  the  hind  wing 
this  row  is  continued,  commencing  at  the  outer  angle  and  proceeding 
to  just  basad  of  the  anal  lobe  on  the  inner  margin.  The  marks  com- 
posing this  line  are  rather  displaced  and  irregular  in  size,  the  one  in 
Cux-Cu2  being  the  largest.  Those  on  the  fore  wing  and  one  on  the  costa 
of  the  hind  wing  are  basally  edged  with  dark  brown  or  black.  Fringe 
of  both  wings  white,  rather  dull,  and  basally  slightly  darkened. 

Length  of  fore  wing.   Female,  13.5  mm.  (neoholotype). 

This  species,  heretofore  considered  synonymous  with  dumetorum, 
is  perfectly  valid.  Dumetorum,  as  described  by  Boisduval,1  has  a 
large  fulvous  area  on  the  underside  of  the  fore  wing,  lacking  in  viridis. 
This  character  (the  extent  of  the  green  on  the  underside  of  the  fore 
wing),  which  so  far  as  known  is  very  constant  and  subject  to  almost 
no  individual  variation,2  is  alone  enough  to  raise  Edwards'  name  from 
the  synonymy.  Oberthur's  figure  of  the  type  of  dumetorum.  (loc.  cit.) 
confirms  Boisduval's  description  in  this  respect.  True  dumetorum  has 
been  discussed  further  above. 

Even  with  viridis  and  dumetorum  separated,  the  picture  is  still  any- 
thing but  clear.  In  California,  apparently  right  along  with  viridis, 
occurs  a  very  green  form  with  an  immaculate  underside.  With  only  a 
limited  number  of  specimens  available,  it  has  been  impossible  to  de- 
termine whether  this  is  a  distinct  species  or  merely  an  extreme  of 
viridis.  Either  is  quite  possible,  although  the  latter  is  the  more 
probable. 

1".  .  .  et  le  disque  des  ailes  superieures  est  beaucoup  plus  largement  roussatre,  ce  qui  fait 
que  le  vert  domine  nioius."    (reference  under  dumetorum). 

2There  is  a  certain  amount  of  individual  variation  in  this  character  in  C.  apama  homopler- 
plera,  as  has  already  been  noted,  but  this  is  apparently  an  indication  of  transition  towards 
another  species  (affinis),  and  as  such  is  excusable. 


228  bulletin:  museum  of  comparative  zoology 

Several  specimens  of  Callophrys  have  been  examined  from  the 
collection  of  Mr.  R.  G.  Wind,  having  been  collected  by  him  chiefly  in 
the  Sierras.  These,  while  apparently  close  to  viridis  differ  in  several 
respects  from  it.  These  specimens,  however,  also  differ  considerably 
inter  se,  so  that  in  the  absence  of  genitalic  examination  or  larger  series, 
nothing  further  can  be  said. 

Callophrys  sheridanii  Edwards 

This  species  occupies  a  long,  narrow,  mountainous  strip  from 
Cloudcroft,  New  Mexico  north  as  far  as  Brewster,  Washington.  It  is 
differentiated  into  at  least  two  subspecies,  the  typical  occurring  in 
Wyoming,  Colorado  and  probably  New  Mexico,  while  the  subspecies 
neoperplcxa  ranges  from  Utah  north  to  Washington.  Specimens  from 
the  latter  locality  do  not  seem  exactly  typical  of  neoperplcxa  and 
when  more  specimens  are  available  they  may  be  found  to  belong  to  a 
distinct  race. 

The  chief  character  whereby  this  species  may  be  differentiated  from 
any  other  now  known  is  the  long,  nearly  straight  (usually)  white  line 
on  the  hind  wing  below,  frequently  basally  bordered  with  black.  It  is 
also  one  of  the  smaller  species  of  the  genus. 

Callophrys  sheridanii  sheridanii  Edwards 

Thecla  sheridanii  Edwards,    1877,  in  Carpenter,  Field  and  Forest,  3,  p.  48 

(lapsus  calami). 
Thecla  sheridanii:  Draudt,  1919  in  Seitz,  Macrolep.  World,  5,  p.  763;  id.,  1924, 

p.  1043. 
Callophrys  sheridanii:  Barnes  and  Benjamin,  1923,  Contrib.  Nat.  Hist.  Lep. 

North  America,  5,  p.  66. 

In  the  Carnegie  Museum  is  one  female  labelled  "Bighorn,  Mont." 
(fide  Sweadner,  in  lift.)  here  selected  as  neoholotype.  Holland's  selec- 
tion of  a  lectotype  from  Denver,  Colorado  is  invalid  as  that  is  definitely 
not  the  type  locality. 

Other  localities.  Wyoming:  Teton  Mts.  and  vie.  (A.M.N.H.). 
Colorado:  Chimney  Gulch  (M.C.Z.);  Ft,  Collins;  Denver  (both 
Barnes  and  Benjamin,  p.  66);  Eldora;  Ceal  Creek  (both  in  coll.  P.  S. 
and  C.  L.  Remington).   New  Mexico:  Cloudcroft  (M.C.Z.). 

Male  above  dark  gray  with  a  slight  brownish  tinge.  A  small  oval 
scent-pad  at  the  upper  cell-end.  Fringes  of  both  wings  white,  gray 
basally.  Female  similar,  but  lacking  the  scent-pad. 

Male  below  with  both  wings  slightly  brassy  green,  irrorated  faintly, 


clench:  notes  on  lycaenid  butterflies  229 

especially  on  the  hind  wing,  with  obscure  black  scales.  Fore  wing  with 
inner  margin  gray-brown.  A  post-discal  line  crosses  from  costa  to  CU2 
(occasionally  with  an  inwardly  dislocated  extension  in  (  u2-2A),  white, 
and  basally  narrowly  lined  with  black-brown.  Hind  wing  with  a 
similar  line,  but  heavier)  and  running  from  costa  to  inner  margin  in  a 
nearly  straight  line  (frequently  dislocated,  but  seldom  if  ever  curved). 
A  small,  white  cell-end  spot  is  occasionally  present.  Fringe  as  on 
upper  surface,  but  slightly  greenish  towards  outer  angle.  Female  as 
in  the  male. 

Length  of  fore  wing.    Male,  10-12  mm.;  female,  11  mm. 

The  typical  form  differs  from  neoperplexa  in  the  thicker  white  line 
below,  and  the  more  prominent  basal  black  edging  to  this  line.  Barnes 
and  Benjamin  state  that  a  specimen  from  Cloudcroft,  New  Mexico  is 
closer  to  the  subspecies  neoperplexa.  This  is  decidedly  queer,  if  so, 
and  is  not  borne  out  by  the  single  specimen  from  that  locality  in  the 
M.C.Z.,  which,  although  not  perfectly  typical,  is  close  enough  to 
belong  here  for  the  present. 

Callophrys  sheridanii  neoperplexa  Barnes  and  Benjamin 

Callophrys  sheridanii  race  neoperplexa  Barnes  and  Benjamin,  1923,  Contrib. 
Nat.  Hist.  Lep.  North  America,  5,  p.  67;  Draudt,  1924,  in  Seitz,  Macrolep. 
World,  5,  p.  1043. 

Holotype  and  allotype  from  Eureka,  Utah.  Paratypes  from  Stock- 
ton and  Silver  Lake,  Utah.  Holotype,  allotype  and  2  male,  1  female 
paratypes  in  the  United  States  National  Museum. 

Other  localities.  Montana :  Polaris  (H.K.C.).  Washington :  Brewster 
(H.K.C.).   In  the  M.C.Z.  is  a  series  from  Silver  Lake. 

Its  differences  as  compared  with  typical  sheridanii  have  been 
pointed  out  above.  The  Brewster,  Washington,  specimens  do  not 
agree  perfectly  with  Utah  neoperplexa,  and  may  ultimately  be  found 
racially  separable.  They  are  very  close,  however,  and  for  the  present 
will  be  left  under  that  name.  It  is  possible  that  some  of  the  Sierran 
material  from  California,  mentioned  under  viridis,  will  prove  to  be 
southern  extensions  of  sheridanii  stock,  racially  modified. 

Length  of  fore  wing.    Male,  10-11  mm.;  female,  11-11.5  mm. 

b.  THE  ACASTE  GROUP  OF  THE  GENUS  THECLA 

The  species  treated  here  form  a  more-or-less  closely  interrelated 
group  isolated  by  Draudt  (1919,  in  Seitz,  Macrolep.  World,  5,  p.  762) 
as  the  tailless  section  of  his  amyntor-group,  of  the  genus  Thccla.    No 


230  bulletin:  museum  of  comparative  zoology 

better  classification  can  be  made  with  accuracy  at  present,  since  the 
generic  subdivision  of  the  neotropical  Theclinae  is  a  task  no  one  has 
yet  attempted  with  any  degree  of  completeness1. 

That  the  "tailless  section"  is  here  considered  separately  from  the 
"tailed  section"  is  due  principally  to  a  very  great  dearth  of  material 
of  the  latter.  The  two  sections  are  apparently  very  closely  allied,  at 
least  in  appearance2,  and  are  separable  only  in  minor  characters. 

The  acaste  group3  may  be  distinguished  from  the  "tailed  section" 
in  the  following  external  particulars :  The  anal  angle  of  the  hind  wing  is 
more  produced,  and  the  anal  lobe  is  longer,  more  prominent.  In 
general  the  tail  is  absent  (see  footnote  no.  3)  but  if  it  is  present  it  is 
coarser  (broader)  and  proportionately  shorter  than  those  of  the  other 
section.  Also  on  the  hind  wing,  the  vein-ends  are  here  slightly  more 
tufted.  The  green  below  is  less  uniform  and  usually  less  shining.  The 
general  appearance  of  this  group  is  more  reminiscent  of  CaUophrys,  or 
even,  somehow,  of  Incisalia,  while  the  others  seem  more  typically 
Thecline. 

The  various  members  of  the  acaste  group  are  of  average  Thecline 
size,  ranging  (in  the  length  of  the  fore  wing)  from  12  (female  of  remits) 
to  16  mm. .(female  of  a.  acaste).  The  males  all  have  scent  pads4,  but 
they  are  usually  small  and  almost  unnoticeable,  occasionally  being  of 
the  same  color  as  the  ground  color,  a  rather  unusual  occurrence.  The 
males  are  above  (excepting  marialis)  some  shade  of  blue  or  purple, 
very  metallic,  with  dark  borders  of  width  varying  with  the  species. 
The  females  on  this  surface  are  duller,  the  blue  being  considerably 
less  metallic,  and  more  basally  restricted.  Below  the  sexes  are  similar, 
with  a  rather  similar  basic  pattern :  ground  color  green ;  a  post-discal 
line  (outwardly  white,  basally  dark)  on  both  wings;  a  submarginal 
row  of  red  spots  on  the  hind  wing;  a  marginal  row  of  spots  or  patches, 
or  a  marginal  band,  on  both  wings;  a  basal  quadrate  patch  of  dark 
brown  on  the  hind  wing.    Any  or  all  of  these  may  be  suppressed  or 

:The  members  of  this  section  (perhaps  even  more  sc  than  those  of  the  "tailed  section")  seem 
to  bear  affinity  to  the  genus  CaUophrys  Billberg.  Indeed.  W.  D.  Field  (1939,  Univ.  Kansas 
Sci  nee  Bull.,  26,  p.  347)  has  placed  Thecla  hercdotus  Fabr.  (a  member  of  the  "tailed  section") 
in  this  genus,  without  comment  however. 

2Due  to  the  briefness  of  time  available  for  study  it  was  not  possible  to  make  any  genitalic 
preparations.    These,  when  examined,  may  show  further,  more  real  differences. 

3So  called  in  preference  to  the  "tailless  section"  of  Draudt,  since  that  method  of  division  has 
proved  incorrect.  Thecla  longula  (pastor)  and  Thecla  marialis,  both  tailed  species,  are  clearly 
members  of  the  group  now  under  consideration,  while  a  tailless  species,  quite  obviously  belong- 
ing to  the  other  section,  is  being  described  as  new  in  another  paper. 

4Godman  and  Salvin  (1887,  Biol.  Centr.-Am.  Lep.,  Rhop.  2,  p.  34)  erroneously  characterize 
agricolor  as  lacking  a  scent  pad. 


clench:  notes  on  lycaenid  butterflies  231 

variously  developed.    An  inner  marginal  hand- of  tan  or  gray  is  always 
present  on  the  underside  of  the  fore  wing. 

The  several  species  together  occupy  a  wide  range  from  Mexico  to 
Bolivia  and  southern  Brazil,  and  even  Argentina.  Our  knowledge  of 
the  distribution  of  the  individual  species  is  very  imperfect,  and  will 
probably  remain  so  for  some  time  to  come. 

Key  to  species 

This  key  was  very  difficult  to  compose,  and  in  parts  may  be  in- 
accurate; it  should  be  used,  therefore,  with  this  in  mind.  In  several 
of  the  species  but  one  sex  is  known,  thus  making  it  impossible  to  in- 
clude characters  which  are  definitely  known  to  apply  to  both  male 
and  female  of  such  species. 

1 .  Frons  brown 2 

-  Frons  green 7 

2.  Outer  margin  of  hind  wing  below  with  red-gray  edging,  either  as  a  definite 

band,  or  as  a  series  of  internervular  spots 3 

-  This  margin  without  red-gray  spots,  being  green,  as  in  the  rest  of  the 

wing 6 

3.  Outer  margin  of  fore  wing  below  with  red-gray  edging;  a  well-marked 

discal  line  as  well  on  this  surface a.  agricolor  (1) 

-  This  margin  without  red-gray  edging;  discal  line  obsolescent  or  wanting  .  .4 

4.  Tail  at  Cu2  on  secondary longula  (5) 

-  No  tail  at  Cu2 5 

5.  Small,  expanse  less  than  30  mm.   (usually  about  26  mm.) remits  (2) 

-  Larger,  over  30  mm.;  up  to  32  mm.  or  so agricolor  banosensis  (la) 

6.  Male  with  scent  pad  black;  underside  nearly  uniform  green  .  .longuloides  (4) 

-  Male  with  scent  pad  colored  blue  as  in  the  ground  color;  below  with  bands 

of  darker  green pseudolongula  (3) 

7.  Upperside  of  male  brown;  tailed  at  Cu2  on  secondary marialis  (9) 

-  This  surface  of  male  bright  blue;  no  tail  at  C112 8 

8.  White  discal  line  on  underside  of  hind  wing  (may  tend  toward  obsolescence) 

9 

-  No  white  discal  line  on  this  wing legionis  (6) 

9.  White  patch  in  center  of  gray  area  on  underside  of  fore  wing  (only  in 

male?) portoena  (8) 

-  No  white  patch  in  this  area,  male  or  female 10 

10.  Submarginal  red  spots  on  underside  of  hind  wing  absent  (almost  or  com- 
pletely); white  discal  line  prominent acaste  catharinetisis  (7a) 

—    Submarginal  red  spots  on  underside  of  hind  wing  usually  well-developed; 
discal  white  line  obsolescent a.  acaste  (7) 


232  bulletin:  museum  of  comparative  zoology 

1.   Thecla  agricolor  agricolor  (Butler  and  Druce) 

Strymon  agricolor  Butler  and  Druce,  1872,  Cist.  Ent.,  1,  p.  105;  Butler,  1873, 

Lep.  Exot.,  p.  158,  pi.  57,  fig.  4. 
Thecla  agricolor:  Hewitson,  1877,  111.  Diurn.  Lep.  Lycaenidae,  p.  201;  Godman 

and  Salvin,  1887,  Biol.  Centr.-Am.    Lep.  Rhop.,  2,  p.  34,  pi.  52,  figs.  11, 

12;  Draudt,  1919,  in  Seitz,  Macrolep.  World,  5,  p.  762,  pi.  154a;  Hoffman, 

1940,  An.  Inst,  Biol.  Mex.,  11,  p.  707. 

The  type  is  in  the  British  Museum,  presumably.  It  was  taken  in 
Cartago,  Costa  Rica.  Draudt  gives  the  range  as  "Mexico  to  Panama." 
Godman  and  Salvin  list  Jalapa,  Mexico;  Duenas,  Guatemala;  Irazu 
and  Rio  Sucio1,  Costa  Rica;  Bugaba  and  Chiriqui,  Panama.  Hoffman 
gives  "Tierra  templada  de  Vera  Cruz.  Sur  de  Puebla.  Morelos. 
Valle  de  Mexico  (2250  M.)."  Specimens  in  the  M.C.Z.  are  all  from 
Jalapa,  Mexico. 

Above,  in  the  male,  metallic  blue,  duller  than  in  the  other  species. 
Both  wings  with  a  rather  indefinite  and  heavy  dark  border  on  the  outer 
margin.  Costa  of  fore  wing  narrowly  black.  Costa  and  inner  margin 
of  hind  wing  narrowly  gray.  Anal  lobe  of  hind  wing  rusty,  black- 
fringed.  The  scent-pad  is  very  small,  almost  unnoticeable  (see  foot- 
note, p.  230),  and  lies  just  beyond  the  upper  cell-end. 

Below,  fore  wing  gray  from  inner  margin  to  M3,  darker  adjacent 
to  the  cell.  Remaining  area  green.  Outer  margin  from  apex  to  inner 
angle  with  a  moderately  heavy  grayish  border.  A  discal  line  of 
reddish  crosses  from  costa,  disappearing  shortly  below  M3.  A  faint 
submarginal  line  behaves  similarly.  Hind  wing  with  a  broad  marginal 
border,  outwardly  gray,  basally  reddish.  The  basal  limit  of  this  border 
is,  in  the  M3-CU1  and  2A-inner  margin  interspaces,  orange.  A  discal 
line  of  dark  green  and  reddish  scales  crosses  the  wing  obscurely.  Base 
marked  with  a  large,  quadrate,  almost  black  patch.  Anal  lobe  maroon. 

The  female  is  duller  above  than  the  male,  with  the  blue  more  re- 
stricted, leaving  very  broad  costal  and  outer  marginal  borders  on  the 
fore  wing,  and  a  broad  outer  marginal  border  on  the  hind  wing.  It  is 
almost  exactly  similar  on  this  surface  to  pseudolongula,  but  the  outer 
limit  of  the  blue  is  more  sharply  defined. 

Below,  the  female  is  similar  to  the  male,  but  with  the  markings 
slightly  brighter. 

Length  of  fore  wing.    Male,  12.5-13.5  mm.;  female,  14.5  mm. 

'Possibly  this  refers  to  a  Rio  Sucio  in  northern  Colombia,  emptying;  into  the  Gulf  of  Darien 
just  below  the  south-eastern  tip  of  Panama.  The  specimens  on  which  this  record  is  based  may. 
therefore,  be  transitional  to  bahosensis. 


clench:  notes  on  lycaenid  butterflies  233 

la.   Thecla  agricolor  banosensis,  new  subspecies 

Upperside: 

Female.  Both  wings  bright  metallic  blue.  Fore  wing  with  a  costal 
and  outer  marginal  border  of  black-brown,  covering  the  whole  cell-end- 
to-apex  region,  and  narrowing  towards  the  inner  angle.  Hind  wing 
with  a  gray  costal  border,  becoming  black-brown  on  the  outer  angle 
and  outer  margin.  Inner  margin  gray.  Anal  lobe  rusty,  the  color 
extending  basad  slightly,  and  costad  as  far  as  Cu2.  Fringe  of  both 
wings  black-brown,  paler  between  the  veins. 
Underside: 

Female.  Both  wings  green.  Fore  wing  with  a  paler,  apple-green 
marginal  stripe  and  a  discal  row  of  three  obscure  red-brown  spots 
(Mi-M2-M3-Cui).  Inner  margin  to  Cu2  gray,  black  basad,  next  the 
cell.  Between  the  gray  and  the  green,  in  the  Cui-Cu2  interspace, 
is  a  band  of  fulvous.  Hind  wing  with  a  small  black  basal  patch,  a 
discal  transverse  line  of  gray,  thin  costad  of  Cui,  heavier  thence  to 
inner  margin.  A  submarginal  row  of  obsolescent  red  lunules  from  costa 
to  inner  margin,  replaced  costally  by  dark  green.  On  the  outer  margin 
is  a  band  of  hoary  maroon,  basally  scalloped  from  Mi  to  the  anal 
angle.  Anal  lobe  maroon.  Fringe  of  both  wings  white-gray,  darker 
at  the  vein-ends. 

Length  of  fore  wing.   Female,  15  mm. 

Holotype.  Female,  San  Pablo,  Rio  Pastaza,  vie.  Banos,  Ecuador, 
2200  meters(?),  (Clark-Maclntyre),  ex  coll.  F.  M.  Brown,  in  the 
American  Museum  of  Natural  History. 

Remarks.  This  subspecies  differs  from  typical  agricolor1  in  several 
respects,  namely :  the  absence  of  the  marginal  hoary  band  on  the  fore 
wing  below,  reduction  in  size  of  the  black  basal  patch  of  the  hind  wing 
below,  and  the  reduced  size  of  the  marginal  band  of  the  hind  wing 
below,  and  its  differentiation  into  two  bands.  The  blue  color  above  is 
very  bright,  far  brighter  than  in  any  other  female  of  this  group  thus 
far  examined  (except  for  a  single  female  of  pseudolongula,  of  almost 
the  same  intensity).  This  character,  however,  is  in  all  likelihood  of 
no  significance,  being  merely  an  age  factor. 

This  subspecies,  with  its  reduced  markings  below,  strongly  sug- 
gests a  transition  from  agricolor  to  rcmus,  which,  in  turn,  may  connect 
to  pseudolongula.   Many  more  specimens  are  needed,  however,  before 

'The  Jalapa,  Mexico,  specimens  and  the  holotype  of  bafwsensis  were  compared  with  Butler's 
figure  in  the  Cist.  Eat.  (I.e.)  and  the  former  agreed  almost  perfectly.  They  were  therefore  used 
in  the  following  comparison  as  typical  agricolor,  even  though  not  topotypical. 


234  bulletin:  museum  of  compaeative  zoology 

this  suggestion  can  be  proven  or  denied.   Genitalic  examination  would 
help  considerably. 

2.   Thecla  remus  Hewitson 

Thecla  remus  Hewitson,  186S,  Descr.  Lycaenidae,  p.  34;  1877,  111.  Diurn.  Lep. 

Lycaetiidae,  p.  201,  pi.  80,  figs.  655-656;  Draudt,  1919,  in  Seitz,  Macrolep. 

World,  5,  p.  763,  pi.  154b. 
Thecla  deidamia  Burmeister,  1879,  Atlas  de  la  Descr.  Phys.  Rep.  Argentine,  5, 

pt.  2,  p.  24. 

Described  from  Brazil.  Type  (female)  is  presumably  in  the  British 
Museum,  although  Mr.  Goodson  (who  examined  the  British  Museum's 
specimens  of  the  present  group  for  me)  wasn't  exactly  clear  on  that 
point  in  his  notes.  Draudt  in  Seitz  gives  no  additional  information, 
and  does  not  seem  to  have  known  the  species.  Burmeister  records  it 
(as  deidamia)  from  Las  Conchas,  north  of  Buenos  Aires,  Argentina. 
Four  specimens,  all  females,  in  the  M.C.Z.:  three  from  Blumenau, 
Sta.  Catharina,  Brazil,  and  one,  ex  coll.  J.  Doll,  labelled  "Brazilia." 
One  male,  in  the  American  Museum  of  Natural  History  from  "Massa- 
randuba-Blumenau,  Brazil." 

Male  above  brilliant  iridescent  blue,  violet-tinted  in  some  lights. 
Both  wings  edged  with  black  on  the  outer  margins.  Hind  wing  gray 
on  costa  and  inner  margin.   Anal  lobe  dull  maroon. 

Below,  the  male  is  green  on  both  wings.  Fore  wing  with  gray  on 
inner  margin  to  Cu2,  darkened  basally.  Hind  wing  with  a  marginal 
row  of  hoary,  reddish-gray  spots,  almost  connected.  A  discal  row  of 
obsolescent,  irregular  spots  crosses  the  wing,  between  which  and  the 
marginal  row  of  spots  is  a  suggestion  of  a  row  of  red  dashes,  in  M3- 
CU1-C112.   Anal  lobe  dark  maroon. 

The  female  above  with  the  blue  much  duller  and  more  restricted; 
quite  similar  in  appearance  to  the  females  of  most  of  the  other  species 
of  the  group.  Anal  lobe  rusty.  Below  green,  with  the  inner  margin  of 
the  fore  wing  gray,  darker  basad.  Hind  wing  on  the  outer  margin 
with  a  series  of  internervular,  almost  round,  reddish  gray  spots.  This 
wing  crossed  in  the  disk  by  a  rather  tortuous  line  of  white,  interrupted 
frequently,  and  of  varying  intensity,  basally  edged  with  blackish. 
Between  this  line  and  the  marginal  spots  are  one  or  two,  rarely  more, 
red  dashes,  slightly  crescentiform.  Anal  lobe  dark  maroon.  Hewit- 
son's  descriptions  (1868, 1877)  of  the  female  tallies  quite  well  with  what 
is  here  regarded  as  remus,  save  for  a  few  minor  differences.  The  fulvous 
patch  at  the  outer  angle  of  the  hind  wing  below  mentioned  by  Hewit- 


clench:  notes  on  lycaenid  butterflies  235 

son  is  very  likely  the  result  of  wear.  He  mentions  in  both  descriptions 
a  band  of  three  or  four  spots  on  the  fore  wing  below,  near  the  eosta. 
No  indication  of  such  spots  was  found  on  the  four  specimens  examined. 
He  does  not  mention  the  subminimal  scries  of  three  or  four  crescenti- 
form  dashes  on  the  hind  wing.  This  character,  however,  is  apparently 
variable,  being  lacking  in  one  of  the  four  M.(  \Z.  specimens,  and  almost 
lacking  in  another.  On  the  strength  of  these  apparent  differences  it 
was  decided  not  to  select  a  type  locality  for  remus  until  either  speci- 
mens are  found  that  match  more  closely  the  description,  or  it  is 
proven  that  he  had  a  slightly  aberrant  example. 

The  male  agrees  quite  well  with  Hewitson's  1877  description  and 
figure,  save  for  the  "green  tint"  lie  speaks  of.  This,  however,  is  fre- 
quently caused  by  chemical  action,  and  can  be  overlooked.  It  is  above 
close  to  males  of  pseudolongula,  but  may  be  separated  by  the  broader, 
more  apically  thickened  outer  marginal  border. 

Thecla  deidamia  is  a  name  that  ever  since  its  publication  in  1S79 
appears  to  have  been  almost  overlooked.  Seitz  made  no  mention  of  it, 
and  the  only  published  reference  of  it  known  to  the  author  is  that  of 
Weeks  (see  synonymy  under  portocna). 

Weeks  referred  his  name  to  "Ruschew.",  but  this  is  in  error,  as  a 
glance  at  the  original  description  will  show.  Ruscheweyh  collected  the 
specimens,  noted  that  they  were  probably  new,  and  suggested  the 
name  deidamia  (in  Hit  to  Burmeister,  apparently).  Burmeister,  how- 
ever, wrote  the  description  and  applied  the  name  to  it,  wherefore  it 
must  go  to  him. 

With  regard  to  the  present  placing  of  the  name,  it  can  be  said  only 
that  a  comparison  of  Burmeister's  description  with  Hewitson's 
descriptions  and  figures  (1868  and  1877)  and  several  specimens  of 
remus  revealed  an  almost  perfect  resemblance.  Burmeister's  locality 
(Las  Conchas,  nr.  Buenos  Aires,  Argentina)  is  quite  in  accord  with  our 
present  knowledge  of  the  distribution  of  remus,  although  extending 
its  range  somewhat  southward. 

3.   Thecla  pseudolongula,  new  species 

Thecla  longula:  Hewitson,  1877,  111.  Diurn.  Lep.  Lycaenidae,  p.  200,  pi.  80, 
figs.  651,  652,  653,  654;  Draudt,  1919,  in  Seitz,  Macrolep.  World,  5,  p.  762, 
pi.  154a.    {nee  T.  longula  Hewitson,  1868  (q.v.)). 

Eyes  narrowly  ringed  with  white.  Frons  with  two  parallel  rows  of 
long,  partially  erect,  dark  brown  hairs,  flanked  outwardly  (along  the 
rims  of  the  eyes)  with  a  row  of  rusty  scales  on  each  side  and  a  band  of 

/ 


236  bulletin:  museum  of  comparative  zoology 

shorter  hairs,  with  rusty  ones  intermingled,  between  them.  Caudad  of 
the  antennae  is  a  transverse  row  of  long  rusty  hairs.  Collar  of  long 
hairs,  bluish  and  rusty  on  top,  becoming  intermingled  with  white  on 
the  sides.  Palpi  rusty  outwardly,  bluish  white  within;  terminal  joint 
completely  rusty.  Antennae  black  above,  white  annulate  below;  club 
black,  tipped  with  fulvous  and  paler  below  (becoming  white  basad). 
Thorax  of  the  male  above  metallic  blue  or  green,  overlaid  moderately 
with  long,  anally  directed  hairs,  heaviest  next  the  abdomen;  female 
paler,  more  steely  blue  above;  male  below  covered  with  dense  brown 
hair,  paler  in  the  female.  Abdomen  of  the  male  above  metallic  blue, 
very  bright;  belbw,  yellow;  tip  above  and  below  gray:  of  the  female, 
duller,  the  blue  more  steely  and  more  anteriorly  restricted;  below 
similar.  Legs  (absent  in  most  of  the  specimens  examined)  apparently 
brown,  tarsi  black  and  white  annulate. 
Upperside  : 

Male.  Bright  metallic  blue,  greenish  in  some  lights,  purplish  in 
others.  The  scent  pad  is  so  small  as  to  be  almost  unnoticeable,  con- 
sisting merely  of  a  short  row  of  scales  along  the  upper  disco-cellular 
at  the  cell-end.  A  marginal  border,  about  1  mm.  thick,  edges  both 
wings.  On  the  fore  wing  it  is  slightly  thicker  than  on  the  hind  wing, 
and  towards  the  apex  it  expands  still  more  (2-3  mm.).  The  anal  lobe 
is  prominent  and  rust-colored.  Fringe  brown  on  fore  wing;  dull  white 
on  hind  wing,  basally  and  at  the  vein-ends  darker. 

Female.    Dull  gray-blue,  with  very  broad  blackish  borders,  some- 
what narrower  and  basally  crenulate  on  the  hind  wing.  Anal  lobe  as  in 
the  male.   Fringe  also  as  in  the  male. 
Underside: 

Male.  Both  wings  bright  emerald-green,  with  the  inner  margin  of 
the  fore  wing  broadly  gray.  The  hind  wing  is  marked  by  two  trans- 
verse bands  of  lighter  green,  rather  indistinct,  basad  to  the  inner  of 
which  is  a  suggestion  (consisting  usually  of  one  or  two  obscure  points) 
of  a  post-discal  line.  In  the  extreme  base  is  a  smallish  dark  brown  area. 
Anal  lobe  rusty  and  adjoining  it  in  the  CU2-2A  interspace  is  an  obscure 
rusty -hoary  patch. 

Female.   Similar,  but  the  markings  more  distinct. 
Length  of  fore  wing.    Male,  13.5-15  mm.;  female,  14-14.5  mm. 
Holotypc.    Male,  Mapoto,  Ecuador,  ex  A.  G.  Weeks  collection. 
Allotype.   Female,  no  locality,  ex  R.  M.  Gray  collection. 
Paratypes.    On  male,  R.  Guamlo  (or  Guamba-  label  poorly  written), 
Ecuador,  ex  A.  G.  Weeks  collection;  one  male,  "Colombia",  Oct.  10, 
1913,  ex  F.   A.   Eddy  collection;  two  males,  no  locality   (possibly 


clench:  notes  on  lycaenid  butterflies  237 

Bogota,  Colombia),  ex  A.  G.  Weeks  collection;  one  female,  no  Locality 
("So.  Am."),  ex  C.  J.  Paine  collection;  two  males,  one  female,  vie. 
Banos,  Ecuador  (Clark-Maelntyre),  as  follows:  one  male,  Tunguragua, 
1900  meters,  March,  1939;  one  male,  Runtim,  2000-2500  meters, 
Nov.  26,  1938;  one  female,  Rio  Blanco,  1700-1900  meters,  Oct.  19, 
1938. 

Holotype,  allotype,  and  three  male  paratypes,  no.  26223  in  the 
Museum  of  Comparative  Zoology.  One  male  and  one  female  paratype 
in  the  author's  collection.  The  last  three  paratypes  in  the  collection  of 
the  American  Museum  of  Natural  History. 

Remarks.  Quite  different  from  true  longula,  for  comparison  with 
which,  see  under  that  species.  The  closest  ally  of  pseudolongula  yet 
discovered  appears  to  be  longuloides,  from  which  it  differs  in  the 
smaller  scent-pad,  broader  borders  and  larger  anal  lobe.  It  is  also 
allied  to  remits,  but  that  species  has  a  marginal  row  of  hoary  spots  on 
the  hind  wing  below. 

4.   Thecla  longuloides,  new  species 

Eyes  hairy,  narrowly  ringed  with  white.  Frons  consisting  of  two 
parallel  rows  of  long,  erect,  dark  brownish  hairs,  thickly  intermingled 
with  rusty  ones.  Just  outside  these  rows,  paralleling  the  white  eye- 
margin,  is  a  row  of  rusty  scales.  Basad  of  the  antennae  is  a  transverse 
row  of  rusty  hairs,  and  between  the  antennae  a  few  white  ones. 
Collar  of  long,  rusty  and  brown-black  hairs.  Palpi  outwardly  covered 
with  mingled  pale  blue,  rusty  and  gray  scales,  inwardly  almost  en- 
tirely pale-blue.  Antennae  black  above,  white  annulate  below;  club 
black  above,  below  tipped  with  dull  fulvous,  backed  by  white.  Thorax 
of  the  male  above  metallic  greenish  blue,  covered,  chiefly  on  the 
periphery  (behind  the  head,  along  the  sides  above  the  wing  bases  and 
anterior  to  the  abdomen),  with  long,  grayish-rusty  hairs;  thorax  of  the 
female  grayer  blue  above,  without  the  greenish  tinge:  below  tufted 
with  brown  in  the  male,  paler  in  the  female.  Abdomen  of  the  male 
above  brilliant  metallic  green,  below  yellow;  tip  gray  above  and 
below;  abdomen  of  the  female  with  the  blue  more  anteriorly  restricted 
above  (the  remaining  area  gray) ;  below  as  in  the  male.  Legs  black 
and  white  annulate. 
Upperside: 

Male.  Both  wings  brilliant  metallic  blue,  greenish  in  some  lights 
and  purplish  in  others.  Fore  vying  with  an  almost  linear  black  scent- 
pad  lying  along  the  upper  discocellular  at  the  cell-end.    Costa  very 


238  bulletin:  museum  of  comparative  zoology 

narrowly,  outer  margin  slightly  more  broadly  black-bordered.  The 
latter  thickens  apically  and  extends  briefly  basad  on  each  vein.  Hind 
wing  with  costa  and  inner  margin  gray.  Outer  margin  narrowly  black, 
also  extending  basad  for  a  short  distance  on  the  veins.  Anal  lobe  small, 
rusty  colored.  Fringe  of  fore  wing  brown,  paler  outwardly;  of  hind 
wing  similar,  but  whitish  outwardly  between  the  veins. 

Female.  Both  wings  dark  gray-brown,  costa  and  inner  margin  of 
hind  wing  pale  brown.  Fore  wing  with  dull  steely  blue  from  inner 
margin  to  upper  discocellular,  from  base  to  cell-end,  outwardly  down 
to  inner  margin  three-quarters  out.  Hind  wing  similar,  leaving  only 
a  narrow  dark  border  on  the  outer  margin,  hazy  and  indistinct  basad, 
that  thickens  slightly  towards  the  outer  angle.  Anal  lobe  as  in  the 
male.  Fringe  as  in  the  male,  but  darker. 
Underside: 

Male.  Both  wings  uniform  green.  Fore  wing  with  the  inner  margin 
to  just  over  Cua  gray,  becoming  sharply  black  along  the  lower  disco- 
cellular.  Hind  wing  with  two  very  faint  and  obscure  small  white  spots, 
each  lined  basally  with  a  few  red  scales:  one  post-discal  in  the  CU1-CU2 
interspace;  the  other  submarginal  in  the  2A-3A  interspace,  touching 
2A.  The  faintest  indication  of  a  submarginal  band,  almost  un- 
noticeable,  consists  merely  of  a  very  slight  darkening  of  the  green. 
Fringe  as  on  upper  surface.   Anal  lobe  obscurely  dark  rusty  colored. 

Female.  Similar  to  the  male,  but  lacking  the  black  along  the  lower 
discocellular  of  the  fore  wing,  and  with  the  faint  submarginal  band  of 
the  hind  wing  a  little  more  prominent.  The  green  on  the  outer  margin 
of  the  fore  wing  extends  down  more  into  the  (A12-IA  interspace. 

Length  of  fore  wing.    Male,  14  mm.;  female,  13  mm. 

Holoti/pe.  Male,  Coroico,  Bolivia,  May,  1899  (Wm.  J.  Gerhard), 
ex  A.  G.  Weeks  collection. 

Allotype.  Female,  Chulumani,  Bolivia,  Dec.  12,  1898  (Wm.  J. 
Gerhard),  ex  A.  G.  Weeks  collection. 

Holotype  and  allotype,  no.  26224  in  the  Museum  of  Comparative 
Zoology. 

Remarks.  This  species  is  allied  to  pseudolongula,  but  may  be  dis- 
tinguished from  it,  in  the  male,  by  the  much  narrower  outer  marginal 
border  above,  and  the  presence  of  a  larger,  more  definite  scent-pad. 
Both  sexes  have  a  considerably  reduced  anal  lobe  on  the  hind  wing 
(less  than  half  the  size  of  that  occurring  on  pseudolongula).  Below  the 
green  is  more  uniform  than  in  pseudolongula,  with  the  light  and  dark 
transverse  bands  almost  non-existent.  In  the  female  the  outer  margin 
of  the  fore  wing  does  not  seem  to  be  so  convex  as  in  that  sex  of  pscudo- 


clench:  notes  on  lycaened  butterflies  _'■>'> 

languid,  and  the  blue  appears  to  be  duller,  although  of  about  equal 
extent. 

5.   Thecla  longula  Hewitson 

Thecla  longula  Hewitson,  1868,  Descr.  Lycaenidae,  p.  34.  (nee  longulti,  Hewit- 
son, 1877,  and  others). 

Strymon  pastor  Butler  and  Druce,  1872,  Cist.  Ent.,  1,  p.  105;  Butler,  1873, 
Lep.  Exot.,  p.  157,  pi.  57,  fig.  5;  McDunnough,  1938,  Mem.  S.  Cal.  Acad. 
Sci.,  1,  p.  24  (no.  364). 

Thecla  pastor:  Godman  and  Salvin,  1887,  Biol.  Centr.  Am.  Lep.  Rhop.,  2,  p.  34, 
pi.  52,  figs.  8,  9,  10;  Draudt,  1919,  in  Seitz,  Macrolep.  World,  5,  p.  762, 
pi.  154a;  Hoffman,  1940,  An.  Inst,  Biol.  Mex.,  11,  p.  707. 

The  type  apparently  is  not  in  the  British  Museum.  Mr.  F.  W. 
Goodson,  at  Tring,  informs  me  through  Dr.  Riley  that  there  is  no 
specimen  of  longula  (by  which  he  meant  pseudolongula,  probably 
exclusively)  in  the  British  Museum  from  Central  America.  The  four 
specimens  of  ''longula"  from  the  Hewitson  collection  are  all  referable 
to  pseudolongula. 

Five  specimens  of  longula  have  been  examined  and  compared  with 
the  description.  They  seem  typical,  and  in  the  absence  of  a  type  are 
made  neotypes,  as  follows: 

Xcoholotypc.  Male,  Orizaba,  Mexico,  June  1941  (Stallings  and 
Turner). 

Neoallotype.    Female,  same  data  as  above. 

Neoparatypes.  One  female,  same  data  as  above;  two  females, 
Presidio,  Mexico,  June  1941  (Stallings  and  Turner). 

Xeoholotype  and  Neoallotype  deposited  in  the  Museum  of  Com- 
parative Zoology.  One  neoparatype  in  the  collection  of  the  author. 
The  remaining  returned  to  the  collection  of  Mr.  D.  B.  Stallings  and 
Dr.  J.  R.  Turner. 

This  species  has  been  subjected  to  a  rather  peculiar  misidentifica- 
tion,  for  almost  the  whole  of  its  existence  in  the  literature  to  date. 
Hewitson's  original  description  of  longula  is  of  a  totally  different 
insect  from  that  of  his  description  and  figure  of  1877  (in  the  "Illustra- 
tions"), as  can  be  seen  from  the  following  extract  from  it  (1868): 
"Underside  dull  green,  tinted  with  orange  at  the  apex  of  the  anterior 
wing1.    Posterior  wing  without  tails:2  crossed  beyond  the  middle  by 

UVohably  due  to  wear:  see  under  remus. 

2Evidently  variable  in  this  species.  One  of  Godman  and  Salvia's  illustrations  of  pastor 
showed  tailless,  and  one  of  the  type  ineotype<  serii>s  of  bmijula  is  also  naturally  without  tails. 
Possibly  the  species  is  in  a  state  of  either  losing  or  acquiring  them. 


240  bulletin:  museum  of  comparative  zoology 

two  bands  of  indistinct  distant  red-brown  spots :  a  series  of  marginal 
red-brown  spots,  irrorated  with  white:  the  lobe  red-brown."  Compar- 
ing this  description  with  that  of  his  published  1877  (pseudolongida  of 
this  paper)  shows  immediately  that  the  two  are  not  conspecific.  When 
the  above  description  was  checked  against  pastor,  the  true  identity 
of  the  name  was  shown. 

6.   Thecla  legionis,  new  species 

Eyes  hairy,  ringed  with  white  or  pale  green.  Frons  green.  Collar 
and  palpi  in  the  present  specimen  indiscernible.  Antennae  black  and 
white  annulate,  nearly  all  black  above;  club  black,  tipped  with  fulvous, 
more  extensive  and  white-backed  below.-  Thorax  pale  steely  blue 
above,  fulvous  beneath,  rather  pallid  where  visible.  Abdomen  gray 
above  (pale  bluish  basad),  cream  below. 
Upperside: 

Female.  Both  wings  steely  blue.  Fore  wing  broadly  black  on  costa, 
apex  and  outer  margin.  Hing  wing  more  narrowly  so  on  outer  margin. 
Costa  and  inner  margin  gray.  Anal  lobe  rusty.  Fringe  of  fore  wing 
blackish,  slightly  paler  outwardly;  of  hind  wing  sordid  white,  darker 
at  the  vein-ends. 
Underside: 

Female.  Both  wings  green.  Fore  wing  with  inner  margin  from  Cu2 
gray.  Cu2  and  outer  margin  along  this  gray  are  fulvous.  Base  of  gray, 
adjoining  cell,  darker.  Hind  wing  unmarked  save  by  a  series  of 
obscure  reddish  dashes  in  the  M2-2A  interspaces  slightly  basad  of  the 
submarginal  area,  and  two  almost  unnoticeable  white  costal  spots, 
one  each  in  the  Sc-Rs-Mi  interspaces.  Outward  of  the  reddish  dashes 
the  green  seems  a  little  paler.  Fringe  of  fore  wing  fulvous,  outwardly 
obscured  by  gray.  Of  hind  wing  fulvous,  outwardly  whitish,  darker  at 
the  vein-ends. 

Length  of  fore  wing.    Female,  12.5  mm. 

Holotypc.  Female,  Blumenau,  Sta.  Catharina,  Brazil  (B.  Pohl), 
no.  26225  in  the  collection  of  the  M.C.Z. 

Remarks.  This  species,  unfortunately  represented  by  but  a  single 
female,  appears  to  stand  closest  to  acaste.  It  has  the  green  frons  of 
acaste,  but  the  almost  unproduced  anal  angle  of,  for  example,  remus. 
Below  it  looks  somewhat  similar  to  a  small  acaste  without  a  discal 
white  line  on  the  hind  wing.  Above,  the  blue  is  slightly  paler  than  in 
females  of  acaste,  but  this  may  be  due  to  fading.  Below  legionis 
differs  from  typical  acaste  (which  subspecies  it  most  closely  resembles) 


clench:  notes  on  lycaenid  butterflies  241 

in  the  almost  complete  absence  of  a  discal  white  line,  the  only  indica- 
tion being  two  almost  costal  white  spots,  so  faint  as  to  be  hardly 
discernible.  The  fringe  is  also  more  fulvous  than  in  either  subspecies  of 
acaste.  The  fulvous  edging  of  the  inner  marginal  area  of  the  fore  wing 
below  is  also  absent  in  both  acaste  and  catharinensis,  but  this  character, 
like  the  fulvous  patches  mentioned  by  Hewitson  in  the  descriptions  of 
longula  and  remus,  may  be  due  to  Avear. 

7.   Thecla  acaste  acaste  Prittwitz 

Thecla  acaste  Prittwitz,  1865,  Stett.  Ent.  Zeit,,  26,  p.  31S;  Draudt,  1919,  in 

Seitz,  Macrolep.  World,  5,  p.  763,  pi.  154a. 
Thecla  lycimna  Hewitson,  1868,  Descr.  Lycaenidae,  p.  33;  1877,  111.  Diurn. 

Lep.  Lycaenidae,  p.  203,  pi.  80,  figs.  663,  664,  665. 

The  type  locality  of  acaste  is  Corcovado,  near  Rio  de  Janeiro, 
Brazil.  Under  his  lycimna  Hewitson  (1S77)  merely  gives  "Brazil."1 
Mr.  Goodson  has  examined  the  Hewitson  type  for  me,  and  it  is  ap- 
parently quite  in  accordance  with  Prittwitz'  original  description,  and 
has  the  characters  here  used  to  separate  true  acaste  from  catharinensis. 
Rio  de  Janeiro,  Brazil,  is  here  selected  as  the  type  locality  of  lycimna, 
thus  better  insuring  its  permanent  synonymy.  The  type  of  acaste, 
Mr.  Goodson  suggests,  is  probably  either  in  Munich,  Berlin,  or  Greis- 
swald,  granting  the  collections  at  those  localities  to  be  still  intact. 

Draudt  gives  as  records:  Sao  Paulo,  Santa  Catharina  and  Rio 
Grande  do  Sul.  The  last  two  will  probably  refer  to  catharinensis. 
Seven  examples  in  the  M.C.Z.,  Rio  de  Janeiro  and  Canto  Gallo,  Brazil. 
The  range  of  this,  the  typical  subspecies,  appears  to  be  quite  limited. 

The  male  above  is  generally  similar  to  longuloides,  but  with  the 
scent-pad  concolorous  with  the  ground.  The  ground  color  is  slightly 
duller  than  in  longuloides,  especially  marginally,  and  the  outer  margin 
is  more  broadly  black.  Below  it  is  uniform  green,  with  a  gray  inner 
margin  on  the  fore  wing,  an  obsolescent  white  transverse  line  on  the 
hind  wing  running  from  costa  to  inner  margin,  outward  of  which  is  a 
row  of  tiny  bright  red  spots  or  dashes,  the  most  prominent  in  Cui-Cu2. 

Female  above  similar  to  the  female  of  pseudolongula,  but  with  the 
outer  limits  of  the  blue  less  definite.   Below  as  in  the  male. 

Both  sexes  differ  greatly  from  pseudolongula  and  longuloides  in  the 
possession  of  a  green  frons. 

Length  of  fore  wing.   Male,  15-15.5  mm.;  female,  14-16  mm. 

!The  original  description  (1868)  cited  no  locality  whatsoever. 


242  bulletin:  museum  of  comparative  zoology 

7a.   Thecla  acaste  catharinensis,  new  subspecies 
Thecla  acaste:  Draudt,  1919,  in  Seitz,  Macrolep.  World,  5,  p.  763  (partim. 

Upperside: 

Male.  Both  wings  dully  shining  violet  blue,  deepening  slightly 
towards  the  margin.  Outer  margin  of  both  wings  narrowly  black, 
thickening  slightly  towards  the  apex  on  the  fore  wing.  Costa  of  fore 
wing  also  narrowly  black.  Costa  and  inner  margin  of  hind  wing  gray, 
the  latter  shaded  basally  with  bluish.  On  the  hind  wing  the  anal  lobe 
is  rusty,  fringed  with  black. 

Female.  Both  wings  black-brown  with  the  basal  two-thirds  of  the 
fore  wing  and  the  majority  of  the  hind  wing  dull  steely  blue,  leaving  a 
costal,  broad  apical,  narrower  outer  marginal  dark  border  on  the  fore 
wing,  and  a  gray  costa  and  inner  margin  on  the  hind  wing,  with  a  still 
narrower  dark  outer  marginal  border.  Anal  lobe  as  in  the  male. 
Underside: 

Male.  Both  wings  bright  green.  Fore  wing  inner  margin  gray  to 
Cu2,  the  green  encroaching  only  at  the  outer  margin.  Base  of  this 
gray  area  dark  along  the  lower  Dc.  Hind  wing  with  a  prominent, 
mildly  tortuous  white  line,  basally  and  obscurely  bordered  with  red. 
This  line  commences  two-thirds  out  on  the  costa  and  proceeds  nearly 
straight  to  three-quarters  out  on  inner  margin.  Anal  lobe  black, 
shading  into  deep  red  basad.  A  few  minute  black  scales  in  the  sub- 
marginal  area. 

Female.    Similar  to  the  male. 

Length  of  fore  wing.    Male  and  female,  J4  mm. 

Holotype.    Male,  Santa  Catharina,  Brazil,  ex  A.  G.  Weeks  collection. 

Allotype.   Female,  Blumenau,  Sta.  Catharina,  Brazil  (B.  Pohl). 

Paratype.   One  female,  "Brazilia",  ex  J.  Doll  Collection. 

Holotype  and  allotype,  M.C.Z.  no.  26226.  Paratype  in  the  collec- 
tion of  the  author. 

Remarks.  Differs  from  typical  acaste  in  the  more  prominent  and 
complete  white  line  on  the  under  surface  of  the  secondary,  in  the 
absence  of  the  submarginal  red  dots  or  dashes  on  the  same  wing  below, 
and,  in  the  male,  by  the  reduced  marginal  border  above. 

8.   Thecla  portoena,  new  species 

Thecla  deidamia:  Weeks,  1905,  111.  Diurn.  Lep.  (111.  Unfig'd.  Lep.),  1,  p.  19. 
(nee  deidamia  Burmeister:  see  under  remits.  The  identity  of  the  insect 
Weeks  called  deidamia  (loc.  cit.)  is  determined  by  three  specimens  so 
labelled  in  his  collection,  which  are  now  made  the  types  of  portoena.) 


clench:  notes  on  lycaenid  butterflies  243 

Eyes  hairy,  ringed  with  white.  Frons  green.  Collar  dark  rusty 
above,  shading  to  whitish  below.  Palpi  rusty  pale  gray,  dorsally 
dark  brown.  Antennae  black,  white  annulate  below,  and  very  faintly 
above;  club  black,  tipped  with  dull  fulvous.  Thorax  dull  black, 
covered  with  bluish  hairs,  lightly  on  top,  heavier  laterally  and  next 
the  abdomen;  below  grayish  tan.  Abdomen  above  blue  next  the 
thorax,  gray  thence  to  tip;  below  yellow,  gray  at  the  tip.  Legs  black 
and  white  annulate. 
Upperside: 

Male.  Both  wings  shining  lavender  blue,  deepening  towards  the 
margin.  Fore  wing  with  a  rather  broad,  dark  brown  marginal  border, 
thickening  apically.  Costa  with  a  similar,  but  narrower  border. 
Hind  wing  with  costa  pale  gray.  Inner  margin  gray,  overlaid  basad 
with  bluish  scales.  Basal  area  of  wing  overlaid  with  pale  scales,  giving 
a  rather  hoary  appearance  to  this  region.  Outer  margin  narrowly 
black-brown,  extending  slightly  basad  on  the  veins.  Anal  lobe  rusty, 
black-fringed.  Fringe  of  fore  wing  dark  basally,  paler  outwardly;  of 
hind  wing  dark,  white  outwardly  between  the  veins. 
Underside: 

Male.  Both  wings  uniform  green.  Fore  icing  with  the  inner  margin 
gray-tan,  with  a  white  patch  on  the  center.  The  green  encroaches  on 
this  gray-tan  at  the  outer  margin.  Base  of  this  area  somewhat  dark- 
ened. Hind  wing  with  a  nearly  straight  white  discal  line,  basally 
edged  with  a  few  red  scales,  that  runs  from  the  costa  towards  the 
inner  margin,  but  disappears  at  about  M3  or  Cui.  A  submarginal  series 
of  thin  red  dashes  occupies  the  M3-2  A  interpaces,  and  occasionally  even 
further  costad.  Anal  lobe  deep  red,  almost  maroon,  extending  slightly, 
in  the  form  of  a  compact  small  area  of  reddish  irroration,  into  the 
Cu2-2A  interspace. 

Length  of  fore  wing.   Male,  12-13.5  mm. 

Holotype.  Male,  Cusilluni,  Bolivia,  May,  1899  (Wm.  J.  Gerhard), 
ex  A.  G.  Weeks  collection. 

Paratypes.  Two  males:  one  male,  same  data  as  holotype;  one  male, 
Coroico,  Bolivia,  April  20,  1S99  (Wm.  J.  Gerhard),  ex  A.  G.  Weeks 
collection. 

Holotype  and  one  paratype,  M.C.Z.  no.  26227.  One  paratype  in 
the  author's  collection. 

Remarks.  T.  portocna  may  be  distinguished  from  both  typical 
acaste  and  its  subspecies  cathariensis  as  follows :  the  marginal  border  of 
the  fore  wing  above  (male)  is  broader  than  in  either;  below,  the  white 
line  on  the  hind  wing  disappears  before  reaching  the  inner  margin, 


244  bulletin:  museum  of  comparative  zoology 

while  in  both  acaste  and  a.  catharinensis  it  proceeds  all  the  way;  the 
anal  lobe  is  here  smaller  than  in  either,  and  colored  deep  red,  while  in 
acaste  and  its  subspecies  it  is  black;  the  inner  marginal  area  of  the  fore 
wing  below  is  here  gray-tan,  and  has  a  central  white  patch,  while  in 
acaste  and  catharinensis  it  is  gray  and  has  no  such  patch  (in  fact 
portoena  is  the  only  species  at  present  known  to  the  writer  that 
possesses  such  a  peculiar  pattern  character) ;  T.  portoena  has  the  red 
submarginal  spots  as  in  acaste  (s.s.),  but  they  are  more  linear,  and  seem 
to  extend  further  cost  ad. 

It  is  possible  that  portoena  may  be  only  a  subspecies  of  acaste.  The 
differences  between  them,  however,  are  such  that  two  full  species  seem 
involved,  and  while  at  present  portoena  seems  to  be  a  local  representa- 
tive of  acaste,  further  knowledge  of  the  distributions  of  the  involved 
forms  may  prove  otherwise. 

9.   Thecla  marialis,  new  species 

Eyes  ringed  with  pale  green.  Frons  green.  Palpi  fulvous,  scaled 
outwardly  with  green ;  terminal  point  black.  Collar  above  green,  with 
pale  greenish  and  fulvous  hairs,  shading  to  sordid  gray  on  the  sides. 
Thorax  above  black  with  dull  greenish-gray  hairs  frontad,  laterally, 
and  next  the  abdomen,  all  back-directed;  below  covered  with  pale 
rusty  long  hairs.  Abdomen  black-brown  above,  yellow  below,  dark 
gray  at  the  tip.  Legs  gray,  with  pale  annulations. 
Upperside: 

Male.  Both  wings  black-brown.  Fore  wing  with  the  basal  area  over- 
laid with  dull  olive-green,  extending  marginad  roughly  two-thirds. 
On  the  upper  cell-end  is  an  elongated  black  scent-pad,  rather  small. 
Hind  wing  similar  but  with  the  olive-green  shading  restricted  more 
basad.  Anal  lobe  fulvous,  this  color  extending  basad  along  inner 
margin  for  one-third  its  length,  and  along  outer  margin  almost  to  Cu2. 
Tail  at  Cu2,  very  short,  but  definitely  present.  Fringe  of  both  wings 
gray  with  a  brownish  tinge. 
Underside  : 

Male.  Both  wings  emerald-green.  Fore  wing  with  inner  margin  to 
Cu2  gray,  darkening  to  almost  black,  next  the  cell,  and  to  darker  gray 
on  the  outer  margin.  Hind  wing  with  an  almost  non-existent  indication 
of  a  post-discal  line,  the  only  real  relic  being  a  tiny  white  spot  in 
Cui-Ci^,  capped  by  a  minute  red  bar.  Anal  lobe  dark  maroon,  capped 
thinly  basad  by  a  white  line. 

Length  of  fore  wing.   Male,  13  mm. 


clench:  notes  on  lycaenid  butterflies  215 

Holotype.  Male,  Victoria,  Mexico,  February  7,  1942  (Mrs.  Mary 
Alice  Turner),  no.  26569  in  the  Museum  of  Comparative  Zoology. 

Remarks.  From  the  remaining  species  of  the  group  now  under  con- 
sideration this  species  differs  most  remarkably  in  the  utter  absence  of 
the  brilliant  morpho-blue  that  characterizes  the  males.  Other  dis- 
tinguishing characters  are:  the  C112  tail,  present  only  in  this  species 
and  in  longula;  and  the  spread  of  fulvous  from  the  anal  lobe  above. 
The  other  differences  are  of  less  importance,  but  can  be  noted  by 
referring  to  the  formal  description  above. 

This  species  is  a  striking  parallel  to  Thecla  fusius  Godman  and 
Salvin1,  which  belongs  to  the  "tailed  section"  of  Draudt's  amyntor- 
group.  Fusius  (of  which  also  only  the  male  is  known)  is  uniformly 
brown  above,  but  below  is  described  and  figured  as  being  exactly  simi- 
lar to  hcrodotus  Fabr.2  From  Godman  and  Salvin's  figures  the  follow- 
ing presumably  significant  differences  have  been  noted :  The  hind  wing 
is  longer,  more  produced  anally  (a  character  of  the  acaste  group,  as 
opposed  to  the  remainder  ("tailed  section")  of  the  am  yutor -group); 
there  is  a  very  plain  anal  suffusion  of  fulvous  in  marialis,  of  which  no 
indication  is  given  either  in  Godman  and  Salvin's  description, or  their 
figure;  nor  is  there,  in  marialis  any  indication  of  the  bluish  suffusion  in 
the  base,  as  depicted  in  their  illustration ;  below  there  is  no  white  line 
on  the  hind  wing  of  marialis,  while  the  figure  of  fusius  plainly  shows 
one,  and  in  which  fusius  also  agrees  with  hcrodotus. 

There  seems  little  doubt,  therefore,  that  in  spite  of  the  superficial 
resemblance,  we  are  dealing  with  a  full  species,  belonging  even  to  a 
separate  subgroup. 

The  collector  of  this  subspecies,  Mrs.  Mary  Alice  Turner,  said  that 
she  could  not  recall  taking  the  specimen  itself,  but  remembered  that 
collecting  at  Victoria  wras  done  by  an  irrigation  ditch,  on  low  weeds 
in  open  country  away  from  the  forest. 

The  author  wishes  to  thank  Mr.  Don  B.  Stallings,  of  Caldwell, 
Kansas,  in  whose  collection  the  specimen  formerly  rested,  for  his  kind 
donation  of  it  to  the  Museum  of  Comparative  Zoology. 

1  1877,  Biol.  Centr.  Am.  Lep.  Rhop.,  2,  p.  34,  pi.  52,  fig.  6,  7. 

2  Godman  and  Salvin  state  that  the  pattern  below  of  fusius  is  so  similar  to  herodotus  that, 
in  the  absence  of  females,  they  were  almost  inclined  to  regard  it  as  a  dimorphic  male  of  that 
species. 


Bulletin  of  the  Museum  of  Comparative  Zoology 
AT   HARVARD   COL  LEG  E 

Vol.  XCIV,  No.  7 


THE  SOCIAL  VESPIDAE  OF  THE  GUI  AN  AS,  PARTICU- 
LARLY OF  BRITISH  GUIANA 


By  Joseph  C.  Bequaert 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED   FOR  THE  MUSEUM 

August,  1944 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2, 
3,  4,  5  and  6  have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVL 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
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After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT  HA  R  YARD   CO  L L E G  E 

Vol.  XCIV,  No.  7 


THE  SOCIAL  VESPIDAE  OF  THE  GUIANAS,  PARTICU- 
LARLY OF  BRITISH  GUIANA 


By  Joseph  C.  Bequaert 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED   FOR  THE  MUSEUM 

August,  1944 


Xo.  7 — The  Social  Vespidae  of  the  Guianas,  "particularly 
of  British  Guiana 

(Hymcnoptera) 
By  Joseph  C.  Bequaert 

The  present  comprehensive  account  of  the  social  wasps  (Vespidae) 
includes  keys  for  the  identification  of  the  South  American  genera  and 
of  the  species  known  from  any  of  the  three  Guianas.  This  area,  based 
on  political  boundaries,  may  be  regarded  as  a  fairly  natural  unit, 
although  the  fauna  is  similar  to  that  of  the  adjoining  forests  of  northern 
Brazil  and  eastern  Venezuela.  Its  wasp  fauna  is,  moreover,  rather  uni- 
form and  I  have  no  doubt  that  all  the  species  included  in  this  paper 
will  eventually  be  found  in  British  Guiana. 

For  the  present,  only  the  forms  definitely  known  from  British  Guiana 
are  fully  discussed,  all  the  recorded  localities  of  these  being  listed. 
Social  wasps  live  in  colonies  of  several  or  many  individuals  and  are 
therefore  abundant  enough  to  be  collected  casually.  Their  nests  are 
also  an  added  attraction  to  the  general  collector.  Hence  they  are  much 
better  known  than  most  other  Hymenoptera,  with  the  possible  excep- 
tion of  the  social  bees.  Most  probably  few  South  American  species 
remain  to  be  discovered,  and  this  is  particularly  true  of  the  Guianas, 
one  of  the  parts  of  tropical  America  most  thoroughly  investigated  by 
entomologists. 

The  Guianas  are  unusually  rich  in  social  wasps  and  would  be  ideal 
for  the  study  of  their  fascinating  habits.  I  list  from  the  entire  area 
98  structural  species  (and  32  additional  color  varieties),  belonging  to 
16  genera  (all  those  known  from  South  America,  except  three:  Synoe- 
coides,  Clypearia  and  Protonectarina).  Of  these,  84  species  (and  18 
varieties)  are  definitely  recorded  from  British  Guiana,  25  of  them  (and 
8  varieties)  having  not  yet  been  taken  in  either  Dutch  or  French 
Guiana.  The  wasp  fauna  of  the  British  portion  thus  appears  to  be 
somewhat  better  known  than  that  of  the  others,  the  differences  being 
entirely  due  to  insufficient  collecting,  in  my  opinion. 

The  following  43  species  and  varieties  were  taken  at  Kartabo  within 
the  quarter  square-mile  area  of  jungle  extensively  studied  by  the  New 
York  Zoological  Society's  Department  of  Tropical  Research. 

Polistes  versicolor  var.  vulgaris 
Polistes  testaceicolor 
Polistes  melanosoma 
Polistes  deceptor 
Mischocyttarus  carbonarius 


250  bulletin:  museum  of  comparative  zoology 

Mischocyttarus  collarellus 

Mischocyttarus  duckei 

Mischocyttarus  flavicans 

Mischocyttarus  labiatus 

Mischocyttarus  prominulus 

Mischocyttarus  smithii 

Mischocyttarus  superus 

Polybia  liliacea 

Polybia  striata 

Polybia  jurinei 

Polybia  bifasciata 

Polybia  bifasciata  var.  quadricincta 

Polybia  chrysothorax 

Polybia  micans 

Polybia  rejecta 

Polybia  occidentalis 

Polybia  occidentalis  var.  parvula 

Polybia  bistriata 

Polybia  tinctipennis  var.  nebulosa 

Stelopolybia  cajennensis 

Stelopolybia  pollens 

Stelopolybia  obidcnsis 

Stelopolybia  constructrix 

Stelopolybia  testacea 

Stelopolybia  pallipes  var.  anceps 

Metapolybia  cingulata 

Protopolybia  minutissima 

Protopolybia  mimdissima  var.  binominata 

Protopolybia  pumila 

Apoica  pallida 

Apoica  pallida  var.  arborea 

Apoica  pallida  var.  pollens 

Apoica  pallida  var.  albimacula 

Brachygastra  scutellaris 

Synoeca  surinama 

Synoeca  virginea 

Parachartergus  smithii 

Sixteen  additional  forms  listed  below  were  taken  in  nearby  localities, 
within  a  fifteen-mile  radius  from  Bartiea : 

Pseudochartergus  chartergoides 
Mischocyttarus  foveatus 
Mischocyttarus  heliconius 
Mischocyttarus  lemoulti 


bequaert:  social  vespidae  of  ghe  guianas  251 

Mischocyttarus  oecothrix 

Mischocyttarus  rohindicollis 

Polybia  catillifex 

Polybia  dimidiata 

Polybia  gorytoides 

Polybia  rufitarsis 

Polybia  sericea 

Polybia  singularis 

Stelopolybia  paraensis 

Stelopolybia  angulata 

Stelopolybia  fulvofasciata 

Prot/)polybia  holoxantha 

Parachartcrgus  fulgidipennis  var.  griseus 

The  combined  two  lists  of  59  forms  give  a  fair  idea  of  the  wasps  to 
be  found  in  that  particular  section  of  British  Guiana.  Additions  may 
be  expected,  some  of  the  more  common  species  of  the  coastal  lowlands, 
such  as  Polistes  canadensis  and  Brachygastra  lecheguama,  not  being 
included.  Nevertheless,  it  is  fairly  certain  that  several  of  the  Guiana 
species  do  not  occur  in  the  Bartica  area,  particularly  those  that  have 
been  reported  only  from  the  hilly  or  mountainous  areas  near  the 
Amazon  watershed  (Alt.  Roraima,  etc.). 

Acknowledgments.  The  present  study  was  based  upon  material 
derived  from  many  sources.  Foremost  were  the  collections  made  over 
a  number  of  years  in  the  Kartabo  district  by  Dr.  W.  Beebe  and  his 
associates  at  the  Tropical  Research  Station  of  the  New  York  Zoological 
Society.  Additional  specimens  were  collected  by  B.  E.  Dahlgren, 
C.  Geijskes,  A.  Mackie,  the  late  J.  G.  Myers,  J.  Ogilvie,  O.  W.  Rich- 
ards, Neal  Weber,  the  late  W.  M.  Wheeler,  and  F.  X.  Williams.  I 
have  also  included  Guiana  records  from  the  collections  of  the  American 
Museum  of  Natural  History,  Carnegie  Museum,  Cornell  University 
(Dept.  of  Entomology),  Field  Museum  of  Natural  History,  Museum  of 
Comparative  Zoology  and  United  States  National  Museum. 

References  to  the  original  descriptions  of  most  of  the  species  and 
color  forms,  and  their  synonyms,  mentioned  in  this  paper  will  be 
found  in  Ducke's  Catalogue  of  the  Social  Wasps  of  Brazil  (1918,  Rev. 
Mus.  Paulista,  10,  pp.  313-374). 


252  bulletin:  museum  of  comparative  zoology 

Table  of  Neotropical  Genera  of  Social  Vespidae 
(Based  Mainly  on  Females  and  ^Yorkers) 

1.  Third  and  fourth  segments  of  mid  and  hind  tarsi  asymmetrical, 

with  inner  apical  lobe  much  longer  than  outer  one.    First  ab- 
dominal segment  much  narrowed,  stalk-like.  . .  . Mischocyttarus 
All  segments  of  mid  and  hind  tarsi  symmetrical 2 

2.  Thoracoabdominal  muscle  inserted  in  a  narrow,  slit-like  furrow 

at  the  lower  end  of  the  propodeum.   First  abdominal  segment 

short  or  slightly  elongate,  but  not  stalk-like Polistes 

Thoraco-abdominal  muscle  inserted  in  a  broadly  ovate  furrow  at 
the  lower  end  of  the  propodeum 3 

3.  Ocelli  very  large,  nearly  as  wide  as  basal  diameter  of  flagellum. 

Mesepisternum  completely  divided  into  an  upper  and  a  lower 
plate.  Abdomen  long  and  slender,  but  first  segment  not  stalk- 
like   Apoica 

Ocelli  normal  or  slightly  swollen,  much  less  than  basal  diameter  of 
flagellum.   In  doubtful  cases,  mesepisternum  undivided 4 

4.  Scutellum  very  prominent,  its  vertical  hind  face  forming  an  angle 

with  the  horizontal  anterior  portion,  which  projects  beyond  the 
short,  vertical  postscutellum.    First  abdominal  segment  short 

and  narrowed,  but  not  stalk-like Brachygastra 

Scutellum  and  postscutellum  placed  one  behind  the  other  in  one 
slope  or  convexity;  scutellum  sometimes  slightly  more  raised, 
but  not  projecting  beyond  postscutellum 5 

5.  Hind  margin  of  postscutellum  considerably  extended  as  a  sharp 

angle  or  broad  lobe  into  the  median  concavity  of  the  propo- 
deum   6 

Hind  margin  of  postscutellum  either  straight  or  forming  a  slight 
obtuse  angle  or  a  broad  curve 7 

6.  Clypeus  usually  much  longer  than  wide,  with  nearly  straight  api- 

cal margin.  Outer  orbits  very  narrow.  Postscutellum  divided 
into  a  short,  horizontal  basal  area  and   a   vertically  abrupt 

apical  portion Pseudochartergus 

Clypeus  either  wider  than  long  or  at  most  about  as  long  as  wide, 
more  or  less  produced  medially  at  apex.   Outer  orbits  of  normal 

width.   Postscutellum  forming  a  single  oblique  convexity 

Protopolybia 

7.  Labial  palpi  bearing  a  heavy,  erect,  curved  seta  some  distance 

from  the  tip.  Clypeus  not  longer  than  wide,  more  or  less  pro- 
duced medially  at  apex 8 


bequaert:  social  vespidae  of  the  guianas  253 

Labial  palpi  always  of  4  segments,  without  erect,  heavy  seta 
before  the  tip.   Maxillary  palpi  of  6  segments 10 

8.  Maxillary  palpi  of  5  segments;  labial  palpi  of  3  segments.    Small 

species,  with  stalk-like  first  abdominal  segment Lcipomeles 

Maxillary  palpi  of  6  segments.  First  abdominal  segment  sometimes 
narrower  than  second,  but  rarely  stalk-like 9 

9.  Labial  palpi  of  4  segments Pseudopolybia 

Labial  palpi  of  3  segments Parachartergus 

10.  First  abdominal  tergite  distinctly  divided  into  a  slender  basal 

stalk  and  a  broader  apical  portion  which  is  part  of  the  remaining 
swollen  abdomen.    Outer  orbits  narrow.    Clypeus  longer  than 

wide,  with  nearly  straight  apical  margin Charterginus 

First  abdominal  tergite  either  without  basal  stalk  or,  if  stalked, 
the  remainder  of  the  tergite  is  also  much  narrower  than  the 
succeeding  segments  and  not  part  of  them 11 

1 1 .  Clypeus  much  longer  than  wide,  with  nearly  straight  apical  margin. 

Outer  orbits  very  narrow 12 

Clypeus  either  wider  than  long  or  at  most  as  wide  as  long,  rarely 
slightly  longer  (in  some  males  longer  than  wide) ;  apical  margin 
more  or  less  produced,  sometimes  minutely  bidentate.  Outer 
orbits  normal 13 

12.  Abdomen  subsessile,  the  first  segment  not  at  all  stalk-like.     Dor- 

sum of  thorax  flattened.  Postscutellum  oblique  throughout .... 

Synoecoides 

Abdomen  with  the  first  segment  distinctly  stalk-like.   Dorsum  of 

thorax  not  flattened.    Postscutellum  with  a  narrow  basal  area, 

followed  by  an  abruptly  vertical  portion Clypearia 

13.  Abdomen  subsessile,  the  first  segment  not  stalk-like  but  posteri- 

orly part  of  the  remaining  swollen  abdomen.    Postscutellum 
with  a  narrow  basal  area  slightly  tuberculate  in  the  middle, 

followed  by  an  abruptly  vertical  portion Chartergvs 

First  abdominal  segment  more  or  less  stalk-like,  always  much 
narrower  than  the  succeeding  swollen  segments.  Postscutellum 
not  so  divided 14 

14.  First  abdominal  segment  very  long  and  slender,  nearly  as  long  as 

thorax,  with  prominent  spiracular  tubercles Metapolybia 

First  abdominal  segment  much  shorter  than  thorax;  spiracular 
tubercles  weak 15 

15.  First  abdominal  segment  forming  a  linear  stalk  with  subparallel 

sides,    flattened    above;    second    segment    abruptly    widened. 
Clypeus  with  two  minute  apical  teeth Epipona 


254  bulletin:  museum  of  comparative  zoology 

First  abdominal  segment  always  much  wider  at  apex  than  at  base. 
Clypeus  ending  in  one  tooth  or  obtusely  rounded  off  at  apex.  .  16 

16.  First    abdominal    segment    with    slightly    prominent    spiracular 

tubercles ;  remaining  segments  abruptly  widened  at  base,  sharply 

conical  and  somewhat  compressed  apically Synoeca 

Abdomen  elongate-oval  and  more  or  less  depressed 17 

17.  Clypeus  very  wide,  almost  twice  as  wide  as  long  in  the  female. 

Ocelli  far  apart,  as  far  from  one  another  as  from  the  eyes 

Protonectarina 

Clypeus  usually  as  wide  as  long  or  slightly  longer  than  wide  or 

moderately  wider;  very  rarely  almost  twice  as  wide  as  long,  in 

which  case  the  ocelli  are  close  together 18 

18.  Mesepisternum  undivided Polybia 

Mesepisternum  divided  by  an  oblique  suture  into  an  upper  and  a 

lower  plate Stelopolybia 

The  following  names  proposed  for  supra -specific  groups  of  Neotropi- 
cal social  wasps  are  not  given  generic  status  in  this  paper. 

Agelaia  Lepeletier  (1836),  based  upon  Agelaia  fuscicomis  Lepele- 
tier  (1836),  is  at  present  unrecognized.  The  species  was  described 
without  locality.   I  suggest  that  it  may  have  been  a  Polistes. 

Caba  R.  v.  Ihering  (1904)  =  Brachygastra. 

Chartergcllus  J.  Bequaert  (1938).    Subgenus  of  Parachartergus. 

Clypeo polybia  Brethes  (1923)  was  based  on  a  species  of  Mischocyt- 
tarus. 

Coloboclypeus  Brethes  (1926)  is  at  present  unrecognized.  Perhaps  it 
was  based  upon  a  solitary  vespid. 

Eupolybia  Dalla  Torre "(1904)  =  Polybia. 

Gymno polybia  Ducke  (1914)  =  Stelopolybia.  See  the  discussion  under 
that  genus. 

Hypochartergus  Zavattari  (1906).    Subgenus  of  Charter ginus. 

Megacanthopus  Ducke  (1904).   Subgenus  of  Mischocyttarus. 

Melissaia  Shuckard  (1841)  =  Brachygastra. 

Monacanthocnemis  Ducke  (1905).    Subgenus  of  Mischocyttarus. 

Myrapetra  White  (1841)  =  Polybia. 

Nectarina  Swainson  and  Shuckard  (1940)  =  Brachygastra. 

Nectarinella  J.  Bequaert  (1938).    Subgenus  of  Parachartergus. 

Tatua  H.  de  Saussure  (1854)  =  E pi pona. 

Xanthocaba  "Cameron"  Meade  Waldo  (1914)  =  Pseudo polybia. 


bequaert:  social  vespidae  of  the  guianas  255 

Subfamily  POLISTINAE 

Polistes  Latreille  (1802) 

This  genus  is  represented  in  South  America  by  more  structural 
species  than  in  any  other  part  of  the  World.  Those  known  from  the 
Guianas  or  likely  to  occur  there  may  be  separated  by  the  following  key. 

1 .  Mesopleura  with  not  even  a  trace  of  prepectal  suture 2 

Mesopleura  at  least  with  traces  of  prepectal  suture  (in  most  cases 

strongly  marked) 7 

2.  Body   thickset.     Abdomen   ovate-fusiform,    widest    before   mid- 

length,  more  or  less  depressed  apically;  first  tergite  strongly 
convex  and,  in  profile,  abruptly  sloping  toward  the  base,  seen 

from  above  as  wide  at  apex  as  long  or  wider 3 

Body  slender.  Abdomen  elongate-fusiform,  widest  about  mid- 
length,  more  or  less  compressed  apically ;  first  tergite  moderately 
convex  and,  in  profile,  gradually  sloping  toward  the  base,  seen 
from  above  longer  than  wide  at  apex 4 

3.  Female:  head  much  swollen;  oculo-malar  space  longer  than  one- 

third  of  height  of  eye  seen  in  front;  clypeus  not  or  barely  touch- 
ing eyes.  Male:  clypeus  pentagonal,  with  bluntly  pointed  apex, 
little  or  scarcely  separated  from  lower  inner  orbits  .  .  P.  carnifex 
Female:  Head  moderately  swollen;  oculo-malar  space  at  most  as 
long  as  one-third  of  height  of  eye,  usually  shorter;  clypeus  touch- 
ing inner  orbits  for  a  distance  equalling  one-third  to  one-half  of 
oculo-malar  space.  Male:  clypeus  subquadrate,  with  straight 
or  weakly  curved  anterior  margin,  widely  separated  from  inner 
orbits P.  major 

4.  Propodeum  very  distinctly  or  coarsely  striate 5 

Propodeum  finely  or  obsoletely  striate 6 

5.  Occipital  carina  lacking  over  lower  third  of  outer  orbit,  where  the 

cheek  is  completely  rounded  off  into  the  gula;  upper  part  of 
outer  orbit  with  an  irregular  slight  depression  near  marginal 

carina P.  canadensis 

Occipital  carina  continued  over  lower  third  of  outer  orbit,  though 
much  weaker  than  in  upper  part,  the  cheek  separated  from  the 
gula  by  a  distinct  ridge;  upper  part  of  outer  orbit  without  de- 
pression near  marginal  carina P.  apicalis 

6.  Occipital  carina  strong,  continuing  along  lower  outer  orbit  to  base 

of  mandible.   Humeral  collar  of  pronotum  strongly  raised 

P.  goeldii 


256  bulletin:  museum  of  comparative  zoology 

Occipital  carina  low,  absent  along  lower  fourth  of  outer  orbit, 
where  the  cheek  is  rounded  off  into  the  gula.  Humeral  collar  of 
pronotum  moderately  raised P.  versicolor 

7.  Propodeum  coarsely  striate.    Occipital  carina  absent  along  lower 

half  of  outer  orbit,  where  the  cheek  is  rounded  off  into  the  gula. 
Clypeus  touching  eyes  over  a  short  distance  in  female,  widely 
separated  from  them  in  male.    Mesepisternum  divided  by  an 

oblique  suture  into  an  upper  and  a  lower  plate P.  major 

Propodeum  finely  or  obsoletely  striate.  Occipital  carina  usually 
continuing  along  outer  orbit  to  base  of  mandible 8 

8.  Outer  orbit  much  widened  in  upper  third,  where  the  occipital 

carina  is  strongly  raised  and  wing-like.  Male:  clypeus  longer 
than  wide,  broadly  rounded  off  at  apex;  all  segments  of  flagellum 

longer  than  wide P.  occipitalis 

Occipital  carina  not  strongly  raised  nor  wing-like  at  upper  third  of 
outer  orbit 9 

9.  Head  much  swollen ;  outer  orbit  markedly  wider  than  eye  in  profile ; 

occipital  carina  slightly  sinuate  about  mid-height.  Mesepister- 
num without  oblique  suture  dividing  it  into  an  upper  and  a  lower 

plate 10 

Head  not  swollen;  outer  orbit  at  most  as  wide  as  eye  in  profile; 
if  wider,  mesepisternum  partly  divided  by  an  oblique  suture.  11 

10.  Clypeus  touching  eyes  over  a  distance  equal  to  about  one-third 

of  length  of  oculo-malar  space.  (Head  and  thorax  black;  abdo- 
men red) P.  bicolor 

Clypeus  touching  eyes  over  a  distance  equal  to  a  little  over  half  the 
length  of  oculo-malar  space.    (Entirely  oily  black) .  .  P.  deceptor 

11.  Body   thickset.     Abdomen   ovate-fusiform,   widest   before   mid- 

length,  more  or  less  depressed  apically;  first  tergite  strongly 
convex  and,  in  profile,  abruptly  sloping  toward  base,  seen  from 
above  as  wide  at  apex  as  long  or  wider.    Occipital  carina  usually 

weak  along  lower  outer  orbit  near  base  of  mandible 12 

Body  slender.  Abdomen  elongate-fusiform,  widest  about  mid- 
length,  more  or  less  compressed  apically;  first  tergite  moder- 
ately convex  and,  in  profile,  gradually  sloping  toward  base,  seen 
from  above  longer  than  wide  at  apex.  Occipital  carina  usually 
high  and  ridge-like  along  outer  orbit  to  near  base  of  mandible .  13 

12.  Mesepisternum  with  at  least  a  trace  of  oblique  suture.   Clypeus  of 

both  sexes  touching  eyes  over  a  distance  equal  to  at  most  one- 
third  of  length  of  oculo-malar  space ;  lower  subocular  portion  in 
female  longer  than  upper  interocular  part.  Male :  clypeus  ending 


bequaert:  social  vespidae  of  the  guianas  257 

in  a  bluntly  pointed  apex;  thirteenth  segment  of  antenna  slightly 

curved P.  ruficornis 

Mesepisternum  without  even  a  trace  of  oblique  suture.  Clypeus 
of  both  sexes  touching  eyes  over  a  distance  equal  to  one-half  or 
more  of  oculo-malar  space;  lower  subocular  portion  in  female 
about  as  long  as  upper  interocular  part.  Male:  projecting  apex 
of  clypeus  broadly  rounded  off;  thirteenth  segment  of  antenna 
straight P.  pacificus 

13.  Occipital  carina  very  faint  or  obsolete  over  lower  third  of  outer 

orbit.  Face  of  male  much  lengthened;  clypeus  higher  than  wide, 
touching  eyes  over  a  distance  equal  to  about  one-third  of  length 

of  oculo-malar  space.  Thorax  densely  silky P.  svbsericeus 

Occipital  carina  sharp  as  far  down  as  base  of  mandible.  Face  of 
male  not  conspicuously  lengthened;  clypeus  about  as  wride  as 
high.  Thorax  not  densely  silky 14 

14.  Male:  Occipital  carina  very  high  and  collar-like  along  outer  orbit; 

outer  orbit  narrower  than  eye  in  profile ;  clypeus  about  as  wide 
as  long;  mesepisternum  with  a  complete  but  very  weak  oblique 
suture.    (Head  and  thorax  black;  abdomen  red.    Female  not 

seen) P.  erythrogaster 

Occipital  carina  low,  not  collar-like  along  outer  orbit 15 

15.  Outer  orbit  wider  than  eye  in  profile.  Clypeus  of  both  sexes  touch- 

ing eyes  over  a  distance  less  than  length  of  oculo-malar  space. 
Oblique  suture  of  mesepisternum  distinct  in  lower  half,  obsolete 
in  upper  half.  (Head,  thorax  and  two  basal  segments  of  abdo- 
men mostly  russet  with  yellow  markings,  remainder  of  abdomen 

black) P.  testaceicolor 

Outer  orbit  as  wide  as  or  slightly  narrower  than  eye  in  profile. 
Clypeus  of  both  sexes  touching  eyes  over  a  distance  about  equal 
to  length  of  oculo-malar  space.  Oblique  suture  of  mesepisternum 

lacking  or  very  faintly  indicated.    (Almost  entirely  black) 

P.  melanosoma 

Polistes  bicolor  Lepeletier  (1836)  is  known  from  French  Guiana, 
Dutch  Guiana,  Brazil  (Amazon  Basin),  Colombia  and  Peru. 

Polistes  pacificus  Fabricius  (1804).  No  doubt  some  of  the  color 
forms  of  this  species  occur  in  British  Guiana.  The  typical  form  is 
known  from  French  Guiana,  as  well  as  from  Mexico,  Colombia,  Trini- 
dad, Brazil  and  Peru;  var.  liliaciosus  H.  de  Saussure  (1854),  from 
French  Guiana,  Brazil  and  Peru;  var.  actaeon  Haliday  (1836)  is  defin- 
itely known  from  Brazil  and  Paraguay,  with  one  rather  doubtful 


258  bulletin:  museum  of  comparative  zoology 

record  from  French  Guiana;  and  var.  geminatus  Fox  (1898)  I  have 
seen  from  French  Guiana  and  Brazil. 

Polistes  apicalis  H.  de  Saussure  (1858)  was  described  from  "Guiana". 
There  is  as  yet  no  definite  record  from  British  Guiana.  I  have  seen  it 
from  Guatemala,  Honduras  and  Ecuador. 

Polistes  carnifex  (Fabricius,  1775).  This  species  was  discussed  in  a 
paper  published  in  1936  (Rev.  de  Entomologia,  6,  pp.  376-383)  It 
occurs  from  central  Mexico  to  northern  Argentina  (Misiones)  in  a 
number  of  color  forms,  four  of  which  I  have  distinguished  by  name. 
As  it  is  often  confused  with  Polistes  major  (Palisot  de  Beauvois),  I 
have  included  that  species  also  in  my  key,  although  it  does  not  seem 
to  occur  in  the  Guianas.  Typical  carnifex  has  not  been  reliably  reported 
from  the  Guianas,  but  Polistes  variegata  Lepeletier  (1836),  described 
from  Cayenne,  may  have  been  this  form.  Var.  rufipennis  Latreille 
(1817)  is  known  only  from  Panama,  French  Guiana  (Polistes  chloro- 
stoma  Lepeletier,  1836,  from  Cayenne,  is  a  synonym)  and  Dutch 
Guiana;  and  var.  ochreata  Spinola  (1851),  from  Dutch  Guiana,  Trini- 
dad, Tobago,  Colombia,  and  Bonacca  (off  the  coast  of  Honduras). 
Both  these  color  forms  no  doubt  occur  in  British  Guiana. 

Polistes  versicolor  (Olivier,  1791) 

One  of  the  most  common  social  wasps  of  South  America,  where  it 
varies  extraordinarily  in  color.  In  a  revision  published  in  1934  (Rev. 
de  Entomologia,  4,  pp.  147-157),  I  recognized  nine  varieties  by  name. 
I  have  since  described  two  more  (1940,  Ent.  News,  51,  pp.  81-82). 
Four  of  these  are  known  from  the  Guianas. 

1.  Typical  form.  Only  first  and  second  abdominal  tergites  marked 
with  yellow.  I  have  seen  it  from  Brazil,  northern  Argentina,  Paraguay, 
Costa  Rica,  and  British  Guiana  (Demerara).  It  was  described  from 
French  Guiana. 

2.  Var.  vulgaris  J.  Bequaert  (1934).  The  most  common  form  of  the 
species,  with  yellow  markings  on  at  least  tergites  1  to  3,  often  1  to  6. 
I  have  seen  it  from  British  Guiana  (Kartabo),  Panama,  Colombia, 
Venezuela,  Trinidad,  Dutch  Guiana,  Brazil,  Paraguay,  northern  Ar- 
gentina, Peru,  Bolivia,  and  Ecuador. 

3.  Var.  myops  (Fabricius,  1798).  Only  the  second  tergite  is  marked 
with  yellow  spots.  It  was  originally  described  from  French  Guiana. 
I  have  seen  it  from  Peru.  It  has  been  recorded  also  from  Brazil  and 
(perhaps  by  error)  from  Trinidad  and  Paraguay. 

4.  Var.  kaieteurensis  J.  Bequaert  (1934).   Abdomen  without  yellow 


bequaert:  social  vespidae  of  the  guianas  259 

markings,  but  the  apical  segments  somewhat  orange.    Thorax  with 
many  yellow  markings.   Described  from  British  Guiana:  Kaieteur. 

Polistes  subsericeus  H.  de  Saussure  (1854) 

I  have  seen  a  specimen  from  British  Guiana:  Mt.  Roraima.  The 
species  is  also  known  from  Dutch  Guiana,  Brazil  and  Paraguay. 

Polistes  erythrogaster  Ducke  (1905) 

I  have  seen  one  specimen  from  British  Guiana:  Mt.  Roraima.  The 
species  is  also  known  from  the  Amazon  Basin  in  Brazil. 

Polistes  testaceicolor  J.  Bequaert  (1937) 

Synonym:  Vcspa  analis  Fabrieius,  1798  (not  of  Fabricius,  1775). 
"Apparently  a  common  wasp  in  British  Guiana:  Bartica;  George- 
town; Warina,  N.  W.  District;  Tumatumari,  Potaro  River;  Kaieteur; 
Kartabo;  Mt.  Everard;  Arakaka,  I  have  also  seen  it  from  Dutch 
Guiana,  French  Guiana,  Brazil  (Amazon  Basin),  Bolivia,  Colombia, 
Venezuela,  Peru  and  Costa  Rica. 

This  species  is  often  heavily  stylopized.  One  female  bears  seven 
empty  pupae  of  Strepsiptera,  protruding  from  the  hind  margins  of 
second  (1),  third  (2)  and  fourth  (2)  tergites,  and  of  third  (1)  and 
fourth  (1)  sternites.  One  female  from  Bartica  is  labelled  as  taken  on 
carrion  of  agouti. 

Polistes  ruficornis  H.  de  Saussure  (1853) 

The  typical  form  of  this  species  is  known  only  from  Uruguay,  Para- 
guay and  northern  Argentina.  I  described  the  var.  demeraraensis 
3.  Bequaert  (1937)  from  British  Guiana  (Mahaica  River,  Demerara; 
Georgetown;  Blairmont),  and  I  now  also  refer  to  this  form  the  speci- 
mens from  Brazil  and  Bolivia  which  I  had  called  var.  biglumoides  in 
1937  (Arch.  Inst.  Biol.  Veg.,  Rio  de  Janeiro,  3,  p.  180). 

The  unrecognized  Polistes  guyanensis  Cameron  (1912),  described 
from  Potaro  River,  British  Guiana,  was  most  probably  a  color  form  of 
P.  ruficornis  and  perhaps  merely  a  variant  of  var.  demeraraensis. 

Polistes  occipitalis  Ducke  (1904) 

British  Guiana:  Mt.  Roraima.  Also  known  from  French  Guiana, 
Dutch  Guiana,  Brazil,  Bolivia,  eastern  Peru,  and  Colombia. 


260  bulletin:  museum  of  comparative  zoology 

Polistes  melanosoma  H.  de  Saussure  (1853) 

British  Guiana:  Bartica;  Kartabo;  west  bank  of  Demerara  River. 
Also  known  from  Brazil  and  Paraguay. 

Polistes  deceptor  W.  A.  Schulz  (1905) 

British  Guiana:  Kartabo.  Also  known  from  Dutch  Guiana,  Brazil 
and  Peru. 

Polistes  canadensis  (Linnaeus,  1758) 

This,  the  most  widely  distributed  of  the  American  Polistes,  extends 
from  south  of  the  Great  Lakes  in  the  United  States  to  northern  Pata- 
gonia, and  occurs  also  in  some  of  the  Lesser  Antilles,  but  not  in  the 
Greater  Antilles.  In  a  recent  revision  (1943,  in  process  of  publication), 
I  recognize  19  color  forms;  but  2  only  are  known  thus  far  from  the 
Guianas. 

1.  Typical  canadensis.  Fairly  uniformly  light  russet  to  dark 
mahogany-brown;  head  and  thorax  sometimes  slightly  lighter  than 
abdomen.  No  yellow  markings,  or  sometimes  a  narrow  apical  margin 
on  the  first  tergite.  Wings  either  uniformly  purplish-black  or  dark 
russet,  or  more  or  less  russet  toward  the  tips  and  darker  basally.  This 
typical  form  occurs  in  British  Guiana:  Demerara;  Georgetown;  Mt. 
Roraima.  It  is  found  also  in  French  Guiana  and  Dutch  Guiana  and 
ranges  from  southern  Arizona  to  northern  Argentina  and  Rio  Grande 
do  Sul.   It  does  not  occur  in  Canada. 

2.  Var.  infuscatus  Lepeletier  (1836).  P.  canadensis  amazonicus 
W.  A.  Schulz  (1905)  is  a  synonym.  Like  typical  canadensis,  but  occiput 
and  outer  orbits  more  or  less  extensively  yellow.  Wings  either  uni- 
formly purplish-black  or  slightly  to  extensively  russet  apically  and 
darker  at  the  base.  First  tergite  with  or  without  narrow  apical  yellow 
fascia.  This  form  is  perhaps  more  common  in  British  Guiana  than 
typical  canadensis:  Demerara;  Onverwagt.  I  have  also  seen  it  from 
Brazil  (Amazon  Basin),  Dutch  Guiana,  French  Guiana,  Colombia, 
Panama,  Peru  and  Ecuador. 

Polistes  urceolata  "Klug"  Erichson  (1848,  in  Schromburgk,  Reisen 
in  Britisch  Guiana,  3,  p.  590;  no  sex;  British  Guiana)  was  evidently 
based  upon  some  form  of  P.  canadensis;  but  the  description  is  too  brief 
for  recognition. 


bequaert:  social  vespidae  of  the  guianas  261 

Subfamily  POLYBIINAE 

Mischocyttarus  H.  de  Saussure  (1853) 

Mischocyttarus  is  the  largest  Neotropical  genus  of  social  wasps, 
some  73  structural  species  and  several  color  forms  being  described  to 
date  (1943).  This  number  will  be  more  than  doubled  in  the  near 
future,  as  both  Mr.  O.  \V.  Richards  and  Mr.  J.  F.  Zikan  are  revising 
the  genus  and  intend  to  describe  many  new  forms.  Meanwhile  it  is 
possible  only  to  enumerate  those  known  from  British  Guiana. 

Mischocyttarus  carbonarius  H.  de  Saussure  (1853) 

Megacanthopus  ruficornis  Cameron  (1912)  is  probably  a  synonym. 
British  Guiana:  West  bank  of  Demerara  River;  Kaieteur;  Kartabo. 
Also  known  from  French  Guiana,  Brazil,  Bolivia,  Peru  and  Panama. 

Mischocyttarus  cerberus  Ducke  (1918) 

The  typical  form  is  known  from  Dutch  Guiana  and  Brazil.  The  var. 
acheron  Richards  (1940)  was  described  from  British  Guiana:  Mazaruni 
Settlement. 

Mischocyttarus  collarellus  Richards  (1940) 

Common  in  British  Guiana:  Mazaruni  Settlement;  Cuyuni  River,  3 
miles  from  Kartabo;  Potaro  River,  on  trail  between  Tukeit  and  Kaie- 
teur; Moraballi  Creek,  Essequibo  River;  Kartabo;  Aremu,  Bartica 
District.    Also  known  from  Dutch  Guiana,  Brazil  and  Panama. 

Mischocyttarus  collaris  Ducke  (1904) 

British  Guiana:  Berbice  Savannas;  Courantyne  River;  Demerara. 
Also  known  from  Brazil. 

Mischocyttarus  duckei  R.  du  Buysson  (1909) 
British  Guiana :  Kartabo.  Originally  described  from  French  Guiana. 

Mischocyttarus  flavicans  Fabricius  (1804) 

Megacanthopus  goeldii  Ducke  (1905)  is  a  synonym. 
British    Guiana:    Kartabo;    Mazaruni    River.     Also    known   from 
French  Guiana,  Dutch  Guiana,  Brazil,  Bolivia,  Peru  and  Ecuador. 


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Mischocyttarus  foveatus  Richards  (1940) 

Described  from  British  Guiana:  First  Falls  of  Essequibo  River; 
Issororo,  Northwest  District;  Island  near  Monkey  Jump;  Moraballi 
Creek,  Essequibo  River. 

Mischocyttarus  heliconius  Richards  (1941) 

Described  from  British  Guiana:  Moraballi  Creek,  Essequibo  River; 
Potaro  River,  on  trail  between  Tukeit  and  Kaieteur. 

Mischocyttarus  injucundus  (H.  de  Saussure,  1854) 

The  typical  form  is  known  from  Brazil  and  Colombia.  The  var. 
bimarginatus  (Cameron,  1912)  was  described  from  British  Guiana, 
without  more  definite  locality.  I  have  seen  it  from  Demerara;  Maza- 
runi  River;  Blairmont;  and  Georgetown 

Mischocyttarus  labiatus  (Fabricius,  1804) 

Megacanthopus  atriceps  Cameron  (1912)  is  a  synonym. 

British  Guiana:  Mt.  Roraima;  Rupununi  River;  Kaieteur;  west 
bank  of  Demerara  River;  Kartabo;  Bartica.  Also  known  from  French 
Guiana,  Dutch  Guiana,  Trinidad,  Brazil,  Venezuela,  Paraguay,  Peru, 
Colombia  and  Panama. 

Mischocyttarus  lecointei  (Ducke,  1904) 

British  Guiana :  Tukeit ;  Kaieteur.  Also  known  from  French  Guiana, 
Dutch  Guiana  and  Brazil. 

Mischocyttarus  lemoulti  (R.  du  Buysson,  1909) 

British  Guiana:  Moraballi  Creek,  Essequibo  River.  Originally  de- 
scribed from  French  Guiana. 

Mischocyttarus  metathoracicus  (H.  de  Saussure,  1854) 

British  Guiana:  Essequibo  River;  Oko  River,  a  tributary  of  the 
Cuyuni.  Originally  described  from  French  Guiana;  also  known  from 
Brazil  and  Peru. 


bequaert:  social  vespidae  of  the  guianas  263 

Mischocyttarus  metoecus  Richards  (1940) 

Described  from  British  Guiana:  Mazaruni  Settlement.  Also  known 
from  Dutch  Guiana  and  Brazil. 

Mischocyttarus  oecothrix  Richards  (1940) 

Described  from  British  Guiana:  Kaieteur  Savanna,  Potaro  Valley; 
Canister  Falls,  Cattle  Trail  Survey;  Moraballi  Creek,  Essequibo  River. 

Mischocyttarus  prominulus  Richards  (1941) 

Described  from  British  Guiana:  Kartabo;  Mazaruni  Settlement. 
Also  known  from  Bolivia. 

Mischocyttarus  rotundicollis  (Cameron,  1912) 

Originally  described  from  British  Guiana,  without  more  definite 
locality.  I  have  seen  it  from  Kalacoon  and  Penal  Settlement,  Bartica 
District. 

Mischocyttarus  smithii  H.  de  Saussure  (1853) 

Megacantkopus  violaceipennis  Cameron  (1912)  is  a  synonym. 
British  Guiana :  Kartabo ;  Essequibo  River.  Also  known  from  French 
Guiana  and  Brazil. 

Mischocyttarus  superus  Richards  (1940) 

Described  from  British  Guiana:  Mazaruni  Settlement;  Bartica- 
Potaro  Road;  Potaro  River,  on  trail  between  Tukeit  and  Kaieteur; 
I  have  also  seen  it  from  Kartabo. 

Mischocyttarus  surinamensis  (H.  de  Saussure,  1854) 

British  Guiana:  Kamakusa;  Courantyne  River;  Trail  between 
Tukeit  and  Kaieteur;  Demerara.  Also  known  from  Dutch  Guiana  and 
Brazil. 

Mischocyttarus  synoecus  Richards  (1940) 

Described  from  British  Guiana:  Amatuk,  Potaro  River;  Mazaruni 
Settlement.    Also  known  from  Brazil  and  Panama. 

Polybia  nigriceps  Cameron  (1912),  described  from  British  Guiana, 
without  more  definite  locality,  was  a  species  of  Mischocyttarus.    The 


264  bulletin:  museum  of  comparative  zoology 

name  is  preoccupied  by  Polybia  fastidiosuscula  var.  nigriceps  Zavattari 
(1906). 

Mr.  O.  W.  Richards  (in  litt.)  includes  British  Guiana  in  the  range  of 
Mischocyttarus  drewseni  (de  Saussure,  1857). 

Mischoeyttarus  socialis  (H.  de  Saussure,  1854)  (=  Vespa  atra  Olivier, 
1791;  not  of  Gmelin,  1790)  was  taken  in  French  Guiana,  Cayenne 
being  the  type  locality  of  Olivier's  V.  atra. 

Mischocyttarus  alfkenii  (Ducke,  1904)  I  have  seen  from  Dutch 
Guiana. 

Pseudochartergus  Ducke  (1905) 

I  have  revised  this  genus  in  a  recent  paper  (1938,  Rev.  de  Entomo" 
logia,  9,  pp.  103-105),  where  I  recognized  only  two  species.  A  study 
of  more  extensive  material  in  the  collections  of  Cornell  University  has 
led  me  to  separate  a  third  species,  Pseudochartergus  panamensis  (Zavat- 
tari, 1906)  (  =  Chartergus  acutiscutis  Cameron,  1907).  This  differs  from 
P.  chartergoides  in  the  relatively  shorter  clypeus  of  the  female  (about 
as  wide  as  long)  and  in  the  lateral  ridges  of  the  propodeum  being  more 
rounded  off.  All  localities  from  British  Honduras,  the  Republic  of  Hon- 
duras, Panama  and  Colombia,  which  I  listed  in  1938  under  P.  charter- 
goides var.  cinctellus,  refer  to  P.  panamensis. 

Only  one  of  the  three  species  is  known  from  the  Guianas. 

Pseudochartergus  chartergoides  (Gribodo,  1891) 

I  distinguish  by  name  three  color  forms  of  this  species,  but  only  the 
var.  cinctellus  (Fox,  1898)  occurs  in  British  Guiana:  Kartabo.  It  is 
known  also  from  Dutch  Guiana  and  Brazil. 

Charter  ginus  pallidibalteatus  Cameron  (1912),  described  from 
British  Guiana,  was  most  probably  P.  chartergoides  var.  cinctellus. 

Charterginus  Fox  (1898) 

This  genus  was  revised  in  1938  (Rev.  de  Entomologia,  9,  pp.  99- 
103).   Only  one  of  the  four  species  is  known  from  the  Guianas. 

Charterginus  huberi  Ducke  (1904) 

British  Guiana:  Source  of  Essequibo  River;  Waratuk.  It  was  de- 
scribed from  the  Amazon  Basin  of  Brazil.  I  have  seen  it  also  from 
Dutch   Guiana   and   French   Guiana    (Maroni).     Polybia  fuhicauda 


bequaert:  social  vespidae  of  the  guianas  265 

Cameron   (1912),  described  from  British  Guiana,  is  a  synonym  of 
C.  huberi. 

Protopolybia  Ducke  (1905) 

This  genus,  containing  some  of  the  smallest  South  American  wasps, 
is  well  represented  in  the  Guianas.  In  addition  to  the  species  included 
in  the  key,  three  forms  described  by  Cameron  from  British  Guiana 
are  as  yet  unrecognized. 

Key  to  Guiana  Species 

1.  Concavity  of  propodeum  narrow,   groove-like.    First  abdominal 

tergite  slender,  the  basal  third  or  more  stalk-like.  Clypeus  about 

as  wide  as  high 2 

Concavity  of  propodeum  broad,  either  very  shallow  or  deeply 
bowl-shaped.  First  abdominal  tergite  thickset,  not  or  very  briefly 
stalk-like  at  base 3 

2.  Thorax  rather  dull,  with  distinct,  scattered,  medium-sized  punc- 

tures; abdomen  impunctate.  Postscutellum  nearly  twice  as  wide 
as  greatest  length,  with  triangular,  bluntly  pointed  apical  exten- 
sion. (Extensively  and  fairly  uniformly  testaceous-yellow ;  almost 

bare) P.  holoxantka 

Body  dull  or  slightly  shiny,  usually  with  a  few  discrete  larger 
punctures.  Postscutellum  at  most  one  and  one-half  times  as  wide 
as  greatest  length,  with  the  apical  extension  rounded  off,  tongue- 
like.   (Black,  with  few  or  many  yellowish  markings) 

P.  minutissima 

3.  First  abdominal  tergite  very  broad  and  short,  cap-shaped,  passing 

gradually  into  the  base  of  the  second,  which  is  not  constricted 
from  it.  Outer  orbit  unusually  widened  in  lower  half;  occipital 
carina  low  and  ending  some  distance  from  base  of  mandible. 
Clypeus  nearly  twice  as  wide  as  high.  Thorax  thickset,  about  as 
high  as  long,  with  many  coarse  punctures;  humeral  collar  high, 
shouldered  at  the  sides;  postscutellum  nearly  twice  as  wide  as 
long;  sides  of  propodeum  bulging,  somewhat  compressed,  though 

broadly  rounded  off P.  emortualis 

First  abdominal  tergite  not  cap-shaped,  always  set  off  from  the 
more  swollen  second.  Thorax  longer  than  high;  sides  of  propo- 
deum low,  depressed,  though  broadly  rounded  off 4 

4.  Clypeus  one  and  one-half  times  to  nearly  twice  as  wide  as  high, 

smooth  and  shiny.   Outer  orbit  about  as  wide  as  eye,  not  reced- 


266  bulletin:  museum  of  comparative  zoology 

ing;  occipital  carina  high,  extending  to  near  base  of  mandible. 
Concavity  of  propodeum  deep;  postscutellum  about  one  and  one- 
half  times  as  wide  as  long;  humeral  collar  high,  more  or  less 
shouldered  at  the  sides.  Body  very  shiny,  with  distinct,  scattered, 
medium-sized  to  large  punctures,  with  many  long,  erect  hairs 

P.  picteti 

Clypeus  much  less  than  one  and  one-half  times  as  wide  as  high. 
Outer  orbit  narrower  than  eye,  receding.  Postscutellum  slightly 
though  distinctly  shorter  than  wide.    Body  dull  or  nearly  dull; 

erect  hairs  short  and  sparse 5 

5.  Thorax  about  one  and  one-fourth  times  as  long  as  high.  Body  dull, 
fairly  uniformly  covered  with  small,  discrete  punctures.  Clypeus 
about  one  and  one-third  times  as  wide  as  high.  (Mostly  pale 
yellow) P.  amarella 

Thorax  nearly  one  and  one-half  times  as  long  as  high.   Body  not  or 

slightly  shiny,  with  microscopic,  alutaceous  sculpture  and  a  few 

small,  discrete  punctures.    Clypeus  at  most  one  and  one-fourth 

•  times  as  wide  as  high.    (Black,  marked  with  yellow) .  .P.  pumila 

Protopolybia  picteti  (H.  de  Saussure,  1854)  is  a  widely  distributed 
species,  of  which  I  recognize  by  name  9  color  forms.  Only  one  of  these 
has  been  reported  from  the  Guianas.  P.  picteti  var.  bella  (R.  von  Iher- 
ing,  1903)  was  originally  described  (as  Polybia  bella)  from  Dutch 
Guiana.  It  is  also  known  from  Brazil,  Peru  and  Bolivia.  It  is  char- 
acterized by  the  ground  color  of  the  body  being  black  or  slightly 
brownish-black,  with  many  yellow  markings  on  thorax  and  abdomen 
(usually  no  yellow  stripes  on  mesonotum)  and  3  pale  spots  on  the  base 
of  tergite  2,  the  median  spot  being  transverse  (wider  than  long). 

Protopolybia  holoxantha  (Ducke,  1904) 

This  species  is  probably  rather  common  in  British  Guiana:  Kama- 
kusa;  Moraballi  Creek,  Essequibo  River;  Penal  Settlement,  Bartica 
District.  It  is  also  known  from  French  Guiana  and  Brazil  (Oyapoc 
and  Amazonas). 

Protopolybia  minutissima  (Spinola,  1851) 

I  recognize  four  color  forms  of  this  species  in  the  Guianas. 

1.  Pale  markings  few,  whitish  or  very  pale  yellow;  base  of  tergite  2 
without  cross-band  or  lateral  spots 2 


bequaert:  social  vespidae  of  the  guianas  267 

Pale  markings  more  extensive,  pale  to  bright  yellow;  base  of  tergite 
2  with  a  cross-band  or  a  pair  of  spots 3 

2.  Thorax  not  or  very  little  marked  with  pale  yellow  or  whitish;  no 

spots  on  propodeum typical  minutissima 

Thorax  more  profusely  marked  with  pale  yellow  or  whitish;  a  pair 
of  spots  on  propodeum var.  bitwminata 

3.  Body  profusely  marked  with  yellow;  base  of  tergite  2  with  a  con- 

tinuous and  usually  broad  cross-band var.  sedula 

Body  less  extensively  marked  with  yellow;  base  of  tergite  2  with  a 
pair  of  streaks  or  spots var.  exigua 

The  typical  form  occurs  in  British  Guiana  (Kartabo),  as  well  as  in 
Peru  and  Brazil. 

The  var.  binominata  (YV.  A.  Schulz,  1906)  appears  to  be  the  most 
common  form,  being  known  from  British  Guiana  (Kartabo;  George- 
town; Demerara),  Colombia,  Panama,  Ecuador,  Peru,  and  Bolivia. 
Polybia  minutissima  H.  de  Saussure  (1854)  was  this  form,  not  Spinola's 
typical  minutissima. 

The  var.  exigua  (H.  de  Saussure,  1854)  is  fairly  common  in  Brazil 
and  has  been  reported  from  French  Guiana,  Trinidad,  and  eastern 
Peru  (Iquitos). 

The  var.  sedula  (H.  de  Saussure,  1854)  is  widespread  and  has  been 
taken  in  Dutch  Guiana,  French  Guiana,  Brazil,  Venezuela,  Trinidad, 
Colombia,  Panama  (Darien),  Peru,  Bolivia,  and  Paraguay.  I  regard 
Polybia  diligens  F.  Smith  (1857)  as  a  synonym  of  sedula. 

Protopolybia  amarella,  new  species 

Fig.  1 

Female.  Head  flattened,  slightly  wider  than  thorax;  seen  in  front, 
subcircular,  scarcely  wider  than  high;  from  above,  rectangular  with 
receding  hind  corners,  about  two  and  one-half  times  as  wide  as  long; 
occipital  margin  with  a  broad  and  shallow  inward  curve.  Vertex  and 
genae  margined  by  a  fine  carina,  which  ends  a  short  distance  from  the 
base  of  the  mandible ;  gena  slightly  narrower  than  eye  in  profile.  Oculo- 
malar  space  lacking.  Inner  orbits  about  one  and  one-half  times  as  far 
apart  on  vertex  as  at  clypeus.  Ocelli  fairly  large,  in  a  nearly  equilateral 
triangle ;  posterior  ocelli  slightly  farther  from  eyes  than  from  each  other. 
Interantennal  shield  broad,  not  set  off,  with  a  longitudinal  groove  over 
upper  half;  antennae  somewhat  farther  apart  than  from  eyes.  Frons 
slightly  convex.    Clypeus  at  its  narrowest  about  one  and  one-third 


268 


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times  as  wide  as  high,  irregularly  heptagonal,  contiguous  to  the  eyes 
over  about  one-half  of  sides;  apical  margins  moderately  produced, 
ending  in  a  blunt,  rounded  point.  Mandible  with  an  oblique  cutting 
edge  of  four  sharp,  subequal  teeth.  Antenna:  scape  long,  slender, 
slightly  curved;  second  segment  unusually  long,  only  slightly  shorter 
than  third,  scarcely  swollen;  remainder  of  flagellum  slightly  club- 


Fig.  1.  Protopolybia  amarella  J.  Bequaert.  Female.  A,  body  in  profile;  B, 
first  and  second  tergites  from  above;  C,  postscutellum  and  propodeum  from 
behind;  D,  head  in  front  view. 


shaped.  Thorax  rather  thickset;  in  profile,  about  one  and  one-fourth 
times  as  long  as  high;  from  above,  nearly  one  and  two-thirds  times  as 
long  as  wide  before  tegulae.  Pronotum  nearly  straightly  truncate 
anteriorly;  humeral  margin  with  a  distinct  obtuse  ridge  which  stops  at 
the  sides,  forming  slight,  blunt  humeral  angles;  dorso-lateral  areas 
sloping;  lower  anterior  margin  of  ventro-lateral  areas  with  a  swelling 
indented  by  a  deep  pronotal  fovea.  Mesopleuron  much  swollen,  with- 
out mesepisternal  suture;  deep  mesepimeral  suture  ending  abruptly 


bequaert:  social  vespidae  of  the  guianas  269 

and  far  from  metapleuron.    Upper  sclerite  of  metapleuron  ending 
nearly  square  below,  not  extended  along  meso-metapleural  suture. 
Scutellum  slightly  and  evenly  convex,  with  a  very  weak  median 
impressed  line.   Postscutellum  triangular,  scarcely  swollen,  its  median 
area  about  one  and  one-third  times  as  wide  as  long,  uniformly  sloping 
or  nearly  vertical  throughout,  the  lower  extension  very  long,  with  a 
free,  raised,  tongue-like  apex.  Propodeum  little  swollen,  nearly  vertical 
in  profile;  median  concavity  very  broad  and  shallow.  First  abdominal 
tergite  short  and  cup-shaped,  with  a  very  short,  broad  stalk,  which  is 
not  wing-like  at  the  sides;  seen  from  above,  gradually  widened,  slightly 
shorter  than  wide  at  apex,  about  half  as  wide  at  apex  as  second  seg- 
ment; in  profile,    very  gradually  and  moderately  convex.      Second 
tergite  from  above  wider  than  long,  gradually  widened  at  base ;  in  pro- 
file, moderately  and  evenly  convex.    Apical  margins  of  all  segments 
very  slightly  thickened.   Mid  tibiae  with  two  spurs.   Claws  symmetri- 
cal, unarmed.    Venation:  second  cubital  cell  much  higher  than  wide; 
third  cubital  long,  about  one  and  one-third  times  as  long  on  cubitus 
as  on  radius ;  radial  cell  very  long,  acute ;  basal  vein  ending  in  subcosta 
Hose  to  the  large  stigma. 

Dull;  fairly  uniformly  covered  with  small,  discrete  punctures,  more 
weakly  on  frons;  clypeus  and  mandibles  almost  impunctate.  Pubes- 
cence sparse,  short,  erect,  somewhat  longer  on  propodeum  and  abdo- 
men.   Eyes  bare. 

Pale  yellow,  with  ferruginous  blotches  on  antennae,  tip  of  mandible, 
tibiae,  tarsi  and  under  side  of  abdomen.  A  few  brownish-black  spots 
as  follows :  a  streak  behind  each  posterior  ocellus ;  a  transverse  row  of 
four  spots  on  the  prehumeral  depressed  slope  of  pronotum;  narrow 
sutures  of  mesonotum,  scutellum  and  postscutellum;  a  median  line 
over  anterior  half,  a  line  on  each  side  over  posterior  half,  and  a  pair  of 
elongate  spots  on  disk  of  mesonotum.  Disk  of  second  tergite  with  a 
median  blackish  triangle  showing  through  the  yellow  integument. 
Wings  hyaline,  with  a  yellowish  tinge;  veins  and  stigma  pale  yellowish- 
testaceous. 

Length  (h.+th.+t.  1+2):  5.1  mm.;  of  fore  wing,  5  mm. 
British  Guiana:  Source  of  Essequibo  River,  female  holotype  (J. 
Ogilvie).  —  Colombia:  La  Chorrera,  Rio  Igara-Parana,  Intendencia 
Amazonas,  female  paratype  (J.  C.  Bradley).  —  Holotype  at  Mus. 
Comp.  Zool.,  Cambridge,  Mass.;  paratype  at  Dept.  Ent.,  Cornell 
University. 

The  almost  completely  yellow  body  makes  this  a  striking  insect. 


270  bulletin:  museum  of  comparative  zoology 

Protopolybia  emortualis  (H.  de  Saussure,  1855) 

This  species  occurs  in  two  color  forms. 

1.  Typical  emortualis  is  more  or  less  extensively  ferruginous,  partic- 
ularly on  the  abdomen,  and  has  pale  yellowish  markings.  It  was  de- 
scribed from  Brazil  and  Ducke  reported  it  from  Dutch  Guiana. 

2.  The  var.  duckei  (R.  du  Buysson,  1905)  (  =  Charter -ginus  duckei 
R.  du  Buysson)  differs  only  in  the  ground  color  of  the  body,  including 
the  abdomen,  being  black.  It  was  originally  described  from  Brazil 
(Bahia) ;  but  Ducke  reports  it  from  eastern  Peru  (Iquitos)  and  I  have 
seen  it  from  British  Guiana  (Upper  Essequibo  River).  It  is  doubtful 
whether  this  form  is  worth  separating  from  typical  emortualis. 

Protopolybia  pumila  (H.  de  Saussure,  1863) 

Two  forms  of  this  species  are  known  in  the.Guianas. 

1.  Typical  pumila  is  extensively  marked  with  yellow,  the  meso- 
notum  often  bearing  yellow  stripes  and  the  base  of  tergite  2  a  pair  of 
unconnected  spots.  I  have  seen  it  from  British  Guiana:  Kartabo; 
source  of  Essequibo  River.  It  is  also  known  from  Dutch  Guiana, 
Brazil,  Venezuela,  Colombia,  Peru,  and  Bolivia. 

2.  The  var.  rotimdata  Ducke  (1910)  was  described  (as  a  species) 
from  French  Guiana  and  is  not  known  from  elsewhere.  It  has  few  pale 
markings,  no  stripes  on  mesonotum  nor  spots  at  the  base  of  tergite  2. 

Protopolybia  rufo-ornata  (Cameron,  1912) 

This  was  described,  as  Charter  ginus  rufo-ornatus,  from  British 
Guiana,  without  more  definite  locality.  It  is  at  present  unrecognized. 
No  doubt  it  was  a  Protopolybia,  possibly  a  color  form  of  P.  picteti. 

Protopolybia  sulciscutis  (Cameron,  1912) 

This  also  was  described  merely  from  "British  Guiana,"  as  Polybia 
sulciscutis,  and  is  unrecognized.  The  statement  in  the  description, 
"apex  of  postscutellum  narrowed  to  a  broad,  bluntly  rounded  point," 
seems  to  place  it  in  Protopolybia. 

Protopolybia  nana  (Cameron,  1912) 

This  is  another  unrecognized  species  described  by  Cameron,  as 
Polybia  nana,  from  "British  Guiana."    The  name  conflicts  with  the 


bequaert:  social  vespidae  of  the  guianas  271 

earlier  Polybia  nana  H.  de  Saussure  (1803);  hut,  as  Cameron's  wasp  is 
probably  identical  with  some  other  described  Protopolybia,  it  would  be 
premature  to  rename  it. 

Brachygastra  Perty  (1833) 

This  genus  has  been  generally  called  Ncctarina  Swainson  and 
Shuckard  (1840) ;  but  there  seems  to  be  no  reason  why  the  earlier  name 
Brachygastra  should  not  be  used. 

Key  to  Guiana  Species 

1.  Sides  of  propodeum  compressed  but  not  angular,  forming  evenly 

rounded  curves  in  profile.  Head  (except  clypeus),  thorax  and 
second  abdominal  tergite  with  deep,  large  punctures.  Dorsal 
(horizontal)  area  of  scutellum  four  times  as  wide  as  long  in  the 

middle B.  Scutellaria 

Sides  of  propodeum  compressed  into  prominent,  broad,  blunt  angles. 
Second  abdominal  tergite  with  small  punctures  only 2 

2.  Dorsal  (horizontal)  area  of  scutellum  nearly  flat,  less  *han  three 

times  as  wide  as  long  in  the  middle,  with  the  hind  margin  rather 
sharp  but  slightly  or  scarcely  curved  inward.  Punctures  of  meson- 
otum  fairly  numerous  and  not  or  scarcely  stronger  than  those  of 
the  base  of  second  tergite.    Clypeus  about  twice  as  wide  as 

high B.  lecheguana 

Dorsal  (horizontal)  area  of  scutellum  convex,  three  to  four  times  as 
wide  as  long  in  the  middle,  with  the  hind  margin  blunt  but  deeply 
curved  inward  (scutellum  forming  two  broad  lobes).  Punctures 
of  mesonotum  decidedly  stronger  than  those  of  second  tergite .  .  3 

3.  Body  with  many  erect,  short  hairs.   Punctures  of  frons  and  meso- 

notum more  numerous  and  fairly  close.  Lateral  angles  of  pro- 
podeum higher.    (Usually  with  many  yellow  markings) 

B.  bilineolata 

Body  with  very  few  erect  hairs.  Punctures  of  frons  and  mesonotum 

far  apart.   Lateral  angles  of  propodeum  lower.   (With  few  yellow 

markings,  the  thorax  and  head  almost  entirely  black) 

B.  augusti 

Brachygastra  augusti  (de  Saussure,  1854) 

The  typical  form  of  the  species  has  distinct  yellow  apical  bands  on 
tergites  1  to  6  and  sternites  2  to  6  of  the  abdomen.   It  is  known  from 


272  bulletin:  museum  of  comparative  zoology 

British  Guiana  (source  of  Essequibo  River),  Dutch  Guiana,  French 
Guiana,  Brazil,  Paraguay,  Bolivia,  Peru,  Colombia,  and  Venezuela. 

Brachygastra  lecheguana  (Latreille,  1824) 

This  widely  distributed  and  common  wasp,  known  from  southern 
Arizona  and  southern  Texas  to  northern  Argentina,  was  discussed  in 
a  paper  published  in  Entomologica  Americana  in  1932  (13,  pp.  92- 
112).  The  specimens  from  British  Guiana  (Arakaka),  Dutch  Guiana 
and  French  Guiana  are  of  the  var.  velutina  Spinola  (1841),  which  has 
the  thorax  densely  covered  with  silky,  golden-yellow  hairs. 

Brachygastra  scutellaris  (Fabricius,  1804) 

The  several  color  forms  of  this  species  are  revised  in  Jr.  New  York 
Ent.  Soc,  50,  (1942)  1943,  pp.  306-308,  where  I  recognize  six  of  them 
by  name.  The  three  known  from  the  Guianas  may  be  separated  as 
follows : 

1.  Scutellutn  black;  postscutellum  either  black  or  with  an  anterior 

pale  cross-band.   Abdomen  with  whitish  apical  margins  on  most 

sternites  and  some  of  the  tergites var.  myersi 

Scutellum  and  postscutellum  mostly  or  entirely  yellow 2 

2.  Second  abdominal   tergite  mostly  orange-yellow  or  ferruginous. 

Thorax  with  many  yellow  markings ;  mesonotum  often  with  spots 

or  stripes var.  rufiventris 

Second  abdominal  tergite  with  only  an  apical  yellow  margin.  Yel- 
low of  pronotum  either  lacking  or  restricted  to  the  narrow,  often 
incomplete  humeral  margin typical  scutellaris 

Typical  scutellaris  is  the  most  common  form  in  British  Guiana: 
Forest  Settlement,  Mazaruni  River;  Kartabo;  junction  of  Mazaruni 
and  Essequibo  Rivers.  Ducke  lists  it  from  Bartica,  and  Bodkin  from 
Issororo,  N.  W.  District.  It  is  also  known  from  French  Guiana,  Brazil, 
Bolivia,  eastern  Peru,  Colombia,  and  Panama.  Brachygastra  scutellata 
Spinola  (1851)  is  a  synonym  of  the  typical  form. 

The  var.  rufiventris  (de  Saussure,  1854)  is  reported  from  French 
Guiana,  Brazil  and  Colombia. 

The  var.  myersi  J.  Bequaert  (1943)  was  described  from  British 
Guiana  (Mt.  Roraima)  and  Bolivia. 


bequaert:  social  vespidae  of  the  guianas  273 

Brachygastra  bilineolata  (Spinola,  1841) 

Recently  (1943,  Jr.  New  York  Ent.  Soc,  50,  for  1942,  pp.  303-306) 
I  recognized  five  color  forms  of  this  species,  those  occurring  in  the 
Guianas  being  separated  in  the  following  key: 

1.  Second  tergite  with  only  the  apical  margin  yellow.    Mesonotum 

with  short  yellow  longitudinal  stripes  or  a  pair  of  yellow  spots 
posteriorly,  or  entirely  black.   Wings  with  a  honey-yellow  tinge, 

most  of  the  veins  testaceous typical  bilineolata 

Second  tergite  with  a  broad  or  narrow  discal  yellow  band  in  addi- 
tion to  the  apical  fascia,  or  mostly  yellow.  Mesonotum  always 
with  two  yellow  stripes 2 

2.  Second  tergite  mostly  yellow,  except  for  the  black  base  and  a  more 

or  less  defined  transverse  black  discal  blotch  or  irregular  line. 

Wings  with  a  honey-yellow  tinge var.  Surinam ensis 

Second  tergite  with  two  separate  transverse  yellow  stripes  of  about 
equal  width,  one  apical,  the  other  discal 3 

3.  Wings  extensively  tinged  with  honey-yellow.    Veins  and  stigma 

mostly  pale var.  antillarum 

Wings  nearly  hyaline,  slightly  grayish,  not  suffused  with  yellow. 
Veins  and  stigma  blackish var.  smithii 

Typical  bilineolata  is  known  from  British  Guiana  (Demerara; 
Georgetown),  French  Guiana,  Dutch  Guiana,  Venezuela,  Brazil, 
Colombia,  and  Costa  Rica.  I  regard  Nectarinia  mobiana  de  Saussure 
(1867)  as  a  synonym  of  typical  B.  bilineolata. 

The  var.  antillarum  (Provancher,  1883)  occurs  in  British  Guiana: 
Rapununi  River.  I  have  seen  it  also  from  Dutch  Guiana,  Trinidad, 
Brazil,  eastern  Peru,  and  the  Republic  of  Honduras. 

The  var.  smithii  (de  Saussure,  1854)  has  been  found  in  French 
Guiana,  Dutch  Guiana,  Brazil,  and  Colombia. 

The  var.  surinamensis  J.  Bequaert  (1943)  is  known  only  from  Dutch 
Guiana. 

Chartergus  Lepeletier  (1836) 

As  stated  under  Parachartergus,  the  type  of  this  genus  was  desig- 
nated by  Emile  Blanchard  (1840)  as  Vespa  nidulans  Fabricius  (1793), 
a  synonym  of  Vespa  chartaria  Olivier  (1791).  I  revised  the  genus  in 
1938  (Rev.  de  Entomologia,  9,  pp,  113-115)  under  the  erroneous  name 
Epipona.  I  recognize  two  species,  of  which  only  one  occurs  in  the 
Guianas. 


274  bulletin:  museum  of  comparative  zoology 

Chartergus  chartarius  (Olivier,  1791) 

I  have  seen  this  wasp  from  British  Guiana:  Demerara;  Georgetown. 
It  occurs  also  in  French  Guiana,  Dutch  Guiana,  Brazil,  Paraguay, 
Bolivia,  and  Peru  (Iquitos). 

Synoeca  H.  de  Saussure  (1852) 

Ducke  (1910)  recognizes  only  two  species  of  Synoeca,  but  I  believe 
three  may  be  separated  on  structural  characters,  as  du  Buysson  did 
in  his  Monograph  (1906).  Moreover,  Polistes  virginea  Fabricius 
appears  to  be  the  valid  name  for  the  species  usually  called  S.  irina 
(Spinola). 

1.  Larger;  fore  wing  16  to  18  mm.  long.   Eyes  bare.   Clypeus  smooth 

and  shiny.  Propodeum  either  impunctate  or  with  a  few  minute 
punctures.  Oculo-malar  space  of  worker  and  queen  fully  as  long 
as  sixth  antennal  segment.  Mostly  or  wholly  bluish-black;  wings 

violaceous-black S.  surinama 

Smaller;  fore  wing  13  to  15  mm.  long.  Eyes  with  a  few  short  hairs. 
Clypeus  rather  dull  and  with  scattered  punctures.  Propodeum 
densely  punctate.  Oculo-malar  space  of  worker  and  queen 
shorter  than  sixth  antennal  segment.  Either  bluish-black  or 
partly  ferruginous.   Wings  mostly  yellowish-subhyaline 2 

2.  Thorax  fairly  uniformly  punctate,  the  punctures  of  mesopleura  and 

mesonotum  scarcely  smaller  than  those  of  propodeum.  Scutellum 
convex,  with  very  slight  median  impressed  line.  Swollen  portion 
of  first  abdominal  segment  elongate-triangular  seen  from  above. 
Black  costal  border  sharply  set  off  from  the  remainder  of  the 

wings S.  chalybea 

Punctures  of  mesopleura  and  mesonotum  very  fine,  much  weaker 
than  those  of  propodeum.  Scutellum  gibbose,  with  a  deep  median 
saddle.    Swollen  portion  of  first  abdominal  segment  trapezoidal 

seen  from  above.   Wings  fairly  uniformly  yellowish-russet 

S.  virginea 

Synoeca  surinama  (Linnaeus,  1767) 

A  common  wasp  throughout  Central  and  South  America,  from 
southern  Mexico  to  southern  Brazil.  The  size  of  the  head  varies  con- 
siderably in  this  species  and  seems  to  be  correlated  with  the  total  size 
of  the  specimen.  Perhaps  these  differences  set  off  the  fertile  females 
(queens)  from  the  workers.    Several  other  insects  are  homoeochromic 


bequaert:  social  vespidae  of  the  guianas  275 

with  S.  surinama.  The  most  interesting  of  these  is  the  mantispid, 
Climaciella  chalybea  Erichson,  of  which  I  have  seen  a  specimen  from 
the  Upper  Rio  Huallaga,  Peru.  The  collector  had  evidently  mistaken 
it  for  the  wasp.    Possibly  the  mantispid  lives  in  the  nest  of  Synoeca. 

1.  Typical  form.  Nearly  entirely  bluish-black,  at  most  with  russet 
blotches  on  the  mandibles.  This  is  Vespa  .surinama  Linnaeus  (1767), 
described  from  Surinam,  where  it  is  common.  Vespa  nigricornis 
Olivier  (1791),  Polistcs  cacrulea  Fabricius  (1804)  and  Synoeca  ultra- 
marina  H.  de  Saussure  (1852)  are  synonyms. 

Common  in  British  Guiana:  Arakaka;  Penal  Settlement,  Bartica; 
Rockstone,  Essequibo  River;  -Kartabo;  Georgetown;  Kamakusa; 
Berbice  Savannas;  Blairmont.  I  have  also  seen  it  from  Costa  Rica, 
Colombia,  Venezuela,  Trinidad,  French  Guiana,  Dutch  Guiana,  Brazil, 
Ecuador,  Peru,  and  Bolivia. 

2.  Var.  cyanea  (Fabricius).  The  lower  part  of  the  head  (particularly 
the  clypeus)  is  more  or  less  ferruginous-red;  second  abdominal  tergite 
occasionally  blotched  with  russet.  Specimens  transitional  to  typical 
surinama  are  frequent.  This  is  Vespa  cyanea  Fabricius  (1775),  Synoeca 
azurea  H.  de  Saussure  (1S52)  and  Synoeca  violacca  H.  de  Saussure 
(1852). 

This  form  is  more  widely  distributed  than  typical  surinama  and  is 
known  from  French  Guiana.  I  have  seen  it  also  from  Mexico  (Apat- 
zingan,  State  of  Michoacan),  the  Republic  of  Honduras,  Guatemala, 
Costa  Rica,  Panama,  Colombia,  Brazil,  Paraguay,  Ecuador,  and 
Venezuela. 

Synoeca  virgixea  (Fabricius,  1804) 

This  species  is  rarer  and  not  as  widely  distributed  as  S.  surinama, 
being  apparently  restricted  to  the  northern  half  of  South  America. 
It  is  almost  entirely  ferruginous  or  russet  with  very  slight  or  no  bluish 
sheen.  Fabricius'  description  of  Polistcs  virginea  (1804)  fits  this  wasp 
exactly.  The  statement  "Antennae  nigrae  articulo  secundo  [  =  scape] 
longiore,  testaceo",  agrees  with  the  Synoeca,  not  with  Apoica  pallida. 
Moreover,  W.  A.  Schulz  (1912,  Berlin  Ent.  Zeitschr.,  57,  p.  84)  recog- 
nized Synoeca  irina  in  Fabricius'  type.  Polistcs  irina  Spinola  (1851) 
and  Synoeca  testacea  H.  de  Saussure  (1854)  are  synonyms. 

British  Guiana:  Kartabo;  Junction  of  Mazaruni  and  Essequibo 
Rivers;  Penal  Settlement,  Bartica  District;  Source  of  Essequibo  River. 
I  have  also  seen  it  from  Dutch  Guiana,  Brazil,  Ecuador,  and  Peru. 


276  bulletin:  museum  of  comparative  zoology 

Synoeca  chalybea  H.  de  Saussure  (1852) 

1.  The  typical  form  of  this  species,  as  figured  by  de  Saussure  (1854), 
is  nearly  entirely  bluish-black,  with  or  without  violaceous  reflections ; 
mandibles,  clypeus,  oculo-malar  spaces  and  spots  on  mesopleura  and 
legs  are  yellowish-russet.  The  specific  name  was  originally  spelled 
chalibea,  but  later  (1854)  corrected  to  chalybea  by  the  author  himself. 

British  Guiana:  Shudihar  River.  I  have  also  seen  it  from  Brazil  and 
Peru  and  it  was  originally  described  from  French  Guiana. 

2.  S.  chalybea  var.  splendens  R.  du  Buysson  (1906)  was  originally 
described  as  a  form  of  S.  irina;  but  all  specimens  I  have  seen  have  the 
puncturation  of  the  thorax  and  the  shape  of  the  first  abdominal  seg- 
ment as  in  S.  chalybea.  It  is  more  or  less  extensively  ferruginous  or 
yellowish-russet.  I  have  seen  it  from  Bolivia  and  Peru.  As  Dr.  Wey- 
rauch  collected  both  typical  chalybea  and  var.  splendens  apparently 
from  the  same  nest  at  Satipo,  Peru,  it  is  doubtful  whether  the  name 
splendens  deserves  to  be  retained. 

Metapolybia  Ducke  (1905) 

I  have  reached  the  conclusion  that  there  are  two  structural  species 
in  this  genus,  not  one  as  Ducke  claimed.  They  may  be  separated  as 
follows : 

1.  First   abdominal   segment    slightly  widened    apically  seen    from 
above  and,  in  profile,  very  gradually  thickened  over  posterior 

half.    Ground  color  of  abdomen  reddish-brown M .  suffusa 

First  abdominal  segment  distinctly  widened  apically  seen  from 
above  and,  in  profile,  rather  abruptly  thickened  over  posterior 
third.    Ground  color  of  abdomen  black M.  clngulata 

M.  suffusa  (Fox,  1899)  has  not  yet  been  taken  in  the  Guianas. 
According  to  Fox'  types  from  Corumba,  Brazil,  this  is  the  wasp  with 
the  abdomen  extensively  russet  which  Ducke  (1910)  mentions  from 
Bolivia  and  calls  Metapolybia  pediculata  var.  rufopicta.  I  have  also 
seen  it  from  Peru,  Bolivia  and  Colombia.  At  Restrepo  (Int.  Meta), 
Colombia,  I  found  M.  suffusa  and  clngulata  nesting  side  by  side,  but  in 
distinct  colonies.  They  were  not  in  the  least  aggressive,  although  the 
sting  was  painful. 

Metapolybia  cingulata  (Fabricius,  1804) 

Schulz  (1912)  saw  the  types  of  Eumenes  cingulata  Fabricius  (1804) 
at  Copenhagen  and  recognized  in  them   Polybia  pediculata  H.   de 


bequaert:  social  vespidae  of  the  guianas  277 

Saussure  (1854).  The  extent  of  pale  yellow  markings  varies,  Fabricius' 
cingulata  being  based  upon  specimens  with  only  the  first  and  second 
tergites  margined  with  yellow.  This  is  also  true  of  Gribodo's  Tatua 
decorata  (1896);  while  Polybia  pediculata  var.  unilineata  R.  v.  Ihering 
(1904)  refers  to  specimens  with  only  traces  of  yellow  markings.  There 
is  no  name  available,  and  none  seems  needed,  for  specimens  with  most 
or  all  tergites  margined. 

British  Guiana:  Kartabo;  Bartica.  Also  known  from  Venezuela, 
Trinidad,  Dutch  Guiana,  French  Guiana,  Brazil,  Paraguay,  northern 
Argentina,  Bolivia,  Peru,  Colombia,  Panama,  Costa  Rica,  Guate- 
mala, and  Mexico  (Cordoba,  Vera  Cruz;  Tierra  Colorada,  Guerrero). 

Epipona  Latreille  (1802) 

Emile  Blanchard  in  1840  (Hist.  Nat.  Ins.,  3,  Orth.  Nevr.  Hem.  Hym. 
Lep.  Dipt.,  p.  394)  designated  as  type  of  Epipona,  Vespa  morio 
Fabricius  (1798),  a  synonym  of  Vespa  tatua  Cuvier  (1797).  This  gen- 
eric name  must  therefore  be  used  instead  of  Tatua  de  Saussure  (1854) ; 
while  the  genus  I  called  Epipona  in  1938  should  be  known  as  Charter- 
gus  Lepeletier.  Epipona  was  revised  in  1938  (Rev.  de  Ehtomologia,  9, 
pp.  115-117),  when  I  recognized  two  species,  only  one  of  which  occurs 
in  the  Guianas. 

Epipona  tatua  (Cuvier,  1797)  has  not  yet  been  taken  in  British 
Guiana;  but  it  is  known  from  French  Guiana,  Trinidad,  Venezuela, 
Brazil,  Peru,  Colombia,  Panama,  and  Costa  Rica. 

Polybia  Lepeletier  (1836) 
Key  to  Guiana  Species 

1.  Propodeum  with  more  or  less  distinct  transverse  striation  extend" 

ing  even  over  the  convex  sides.  Occipital  margin  of  head  with  a 

deep  inward  curve.   Body  14  to  16.5  mm.  long 2 

Propodeum  without  or  with  obsolete  striation;  rarely  a  few  irregu- 
lar striae  in  the  concavity  or  on  the  sides 4 

2.  Head  very  strongly  swollen  behind  the  eyes;  outer  orbit  slightly 

wider  than  eye  in  profile.  Clypeus  broader  than  high,  the  apex 
evenly  and  broadly  rounded  off.  Pronotum  on  the  sides  with  a 
strong  but  blunt  humeral  crest.    Striation  of  propodeum  fine, 

but  regular P.  ujhelyii 

Head  moderately  swollen  behind  the  eyes;  outer  orbit  narrower 
than  eye  in  profile.  Clypeus  about  as  high  as  wide,  with  bluntly 
pointed  apex.   Pronotum  somewhat  shouldered,  but  not  ridged 


278  bulletin:  museum  of  comparative  zoology 

at  the  humeral  angles.  Striation  of  propodeum  very  irregular 
or  partly  effaced 3 

3.  Concavity  and  sides  of  propodeum  with  coarse  and  irregular 

wrinkles  and  punctures.  First  abdominal  tergite  slightly  swol- 
len; stalk-like  base  passing  gradually  into  swollen  apical  por- 
tion (in  profile  view) P-  striata 

Striation  and  punctures  of  propodeum  weak  or  obsolete.  First 
abdominal  tergite  strongly  swollen,  abruptly  raised  behind  the 
stalk-like  base  (in  profile  view) P.  liliacea 

4.  Median  concavity  of  propodeum  distinct  and  more  or  less  circular, 

never  longer  than  wide 5 

Median  concavity  of  propodeum  either  almost  lacking  or  groove- 
like and  longer  than  wide 7 

5.  Head  distinctly  swollen;  outer  orbit  fully  as  wide  as  eye  in  profile. 

Puncturation  of  mesopleura  fine  but  distinct.  Black,  with 
scutellum  and  postscutellum  entirely  yellow.    Body  12  to  14 

mm.  long P •  jurinei 

Head  not  swollen;  outer  orbit  narrower  than  eye  in  profile.  Meso- 
pleura nearly  impunctate.   Smaller;  body  8  to  10  mm.  long..  .6 

6.  Pronotum  with  very  weak,  blunt  humeral  collar;  hind  margin 

forming   an   oval   curve.     First   abdominal   segment   slender; 

swollen  portion  longer  than  wide  at  apex P.  signata 

Pronotum  with  distinct,  sharp  humeral  collar;  hind  margin  nearly 
semi-circular.  First  abdominal  segment  shorter;  swollen  portion 
about  as  long  as  wide  at  apex P-  bifasciata 

7.  At  least  sides  of  thorax  covered  with  dense,  silky  pubescence, 

often  with  golden  sheen.    Yellow  markings  lacking  or  much 

reduced.    Body  14  to  19  mm.  long 8 

Thorax  nowhere  covered  with  dense,  silky  pubescence 11 

8.  Eyes  with  distinct  short  hairs.    Oculo-malar  space  very  long  (as 

long  as  fifth  antenna!  segment).  Pronotum  without  humeral 
collar.  Black,  with  thorax  more  or  less  extensively  and  first  or 
first  and  second  abdominal  segments  reddish-brown.    Body  15 

to  17  mm.  long P-  sericea 

Eyes  bare.   Oculo-malar  space  very  short  or  lacking 9 

9.  Dorsum  of  thorax  densely  covered  with  long  golden-silky  pubes- 

cence. Mesopleura  finely  but  distinctly  punctate.  Pronotum 
without  humeral  collar.  Mostly  brownish-red  to  fuscous; 
abdomen  without  paler  apical  margins;  head  black;  wings  more 
or  less  ferruginous,  particularly  toward  costal  margin.  Clypeus 
about  as  long  as  wide.  Body  15  to  17  mm.  long. .  P.  chrysothorax 


bequaert:  social  vespidae  of  the  guianas  279 

Dorsum  of  thorax  only  sparsely  pubescent,  not  golden-silky. 
Color  different 10 

10.  Only  pleura  of  thorax  distinctly  silky;  the  abdomen  not.    Wings 

subhyaline  or  slightly  brownish-yellow,  with  ferruginous  costal 

margin.    Body  15  to  16  mm.  long P.  affinis 

Entire  body,  including  abdomen,  silky  or  velvety.  Either  russet 
to  brownish-russet  with  testaceous  apical  abdominal  margins, 
or  (var.  velutiiia)  almost  entirely  blackish  without  apical  mar- 
gins. Wings  strongly  tinged  with  honey-yellow.  Thorax  very 
finely  punctate.  Pronotum  without  humeral  collar.  Clypeus 
slightly  wider  than  long.    Body  14  to  18  mm.  long. .  .  P.  micans 

11.  Large  species,  18  to  21  mm.  in  total  length.    Head  and  thorax 

black;  abdomen  red;  wings  nearly  hyaline.  Thorax  shiny,  with 
scattered  punctures  and  long,  erect  hairs.    Oculo-malar  space 

long.    Clypeus  much  wider  than  long P.  dimidiata 

Smaller.  Thorax  not  shiny,  or  the  other  characters  mentioned 
above  not  all  present 12 

12.  First  tergite  distinctly  punctate.    Mesopleura  and  propodeum 

with  large  punctures.  Outer  orbits  somewhat  swollen  in  upper 
half.  Clypeus  slightly  wider  than  high.  Eyes  with  distinct  short 
hairs.  Body  black,  somewhat  velvety,  12  to  14  mm.  long;  wings 

infuscate,  clearer  posteriorly  or  apically 13 

First  tergite  usually  impunctate;  if  distinctly  punctate,  the  other 
characters  mentioned  above  not  all  present 14 

13.  Oculo-malar  space  at  its  shortest  over  half  the  length  of  the 

tenth  antennal  segment.  Concavity  of  propodeum  narrow, 
groove-like,  but  very  weak  or  barely  indicated  in  upper  third 

P.  tincti'pennis 
Oculo-malar  space  at  its  shortest  less  than  half  the  length  of  the 
tenth  antennal  segment.    Concavity  of  propodeum  well-devel- 
oped over  entire  length,  broader  and  more  shallow.  .  P.  rufitarsis 

14.  Oculo-malar  space  distinct,  at  least  half  the  length  of  the  tenth 

antennal  segment 15 

Oculo-malar  space  very  short  or  almost  lacking.  Clypeus  always 
much  narrower  than  vertex.  Outer  orbit  usually  narrower  than 
eye  in  profile 17 

15.  Black,  with  partly  black  wings.   Eyes  hairy.   A  short  but  distinct 

carina  between  occiput  and  middle  part  of  outer  orbit  (not 
behind  vortex).   A  fine,  sharp  humeral  collar.  Clypeus  about  as 

high  as  wide P.  ignobilis 

Russet,  infuscate  or  blackish,  with  many  pale  (yellowish)  mark- 


280  bulletin:  museum  of  comparative  zoology 

ings,  particularly  on  the  thorax  (mesonotum  striped).  Wings 
slightly  tinged  with  yellow.  No  carina  between  occiput  and 
outer  orbit  or  vertex.  No  humeral  collar.  Eyes  bare.  Clypeus 
much  wider  than  high.  Outer  orbit  slightly  wider  than  eye  in 
profile 16 

16.  Larger  species;  fore  wing  11  to  11.5  mm.  long.  Oculo-malar  space 

about  as  long  as  tenth  antennal  segment,  longer  than  the  stretch 
of  the  inner  orbit  contiguous  to  the  clypeus.  Clypeus  somewhat 

narrower P.  singularis 

Smaller  species;  fore  wing  8.5  to  9.5  mm.  long.  Oculo-malar  space 
about  half  the  length  of  the  tenth  antennal  segment,  shorter 
than  the  stretch  of  the  inner  orbit  contiguous  to  the  clypeus. 
Clypeus  very  wide ' P.  emaciata 

17.  Large  species  (15  to  18  mm.  long);  black  with  extensive  pale 

markings;  mesonotum  always  with  two  longitudinal  pale  stripes. 
•  Upper  outer  orbit  separated  from  occiput  by  a  strong  carina, 

which  however  does  not  continue  behind  vertex.    Propodeum 

with  at  least  traces  of  striation.    (P.  striata  and  P.  liliacea: 

see  couplet  3) 
If  pale  longitudinal  stripes  are  present  on  mesonotum,  the  species 

is  much  smaller.   Propodeum  without  traces  of  striae 18 

18.  Thorax  shiny,  with  many  distinct  punctures.   A  distinct  but  short 

oculo-malar  space.  Eyes  with  scattered  hairs.  Longitudinal 
groove  of  propodeum  deep.  Russet,  with  yellowish  markings 
and    infuscate    areas.     Wings    nearly    hyaline,    very    slightly 

yellowish P.  gorytoides 

Thorax  practically  impunctate 19 

19.  Humeral  margin  of  pronotum  barely  curved,  raised  on  the  sides 

into  a  sharp,  low  collar  which  projects  as  fairly  distinct  rounded 
angles.  Groove  of  propodeum  weak  or  obsolete ;  propodeum  with 
erect  hairs.    No  carina  between  occiput  and  vertex.    Body  12 

to  13  mm.  long P.  rejecta 

Humeral  margin  of  pronotum  distinctly  curved,  either  not  raised 
or  with  a  low  collar  which  never  forms  angles  at  the  sides ....  20 

20.  Sides  of  pronotum  with  a  distinct  though  low  humeral  collar. 

Upper  outer  orbits  and  vertex  not  separated  by  a  carina  from 
occiput.  Propodeum  distinctly  grooved,  with  erect  hairs.   Body 

9  to  12  mm.  long 21 

Humeral  collar  barely  indicated  or  absent.  In  doubtful  cases,  with 
a  fine  carina  between  occiput  and  vertex 22 

21.  Thorax  slender,  flattened  dorsally,  particularly  on  posterior  por- 


bequaert:  social  vespidae  of  the  guianas  281 

tion  of  mesonotum  and  on  scutellum.    Scutellum  not  grooved. 

Eyes  with  fine,  scattered  hairs P.  procellosa 

Thorax  more  thickset,  not  flattened  dorsally;  scutellum  convex, 

with  a  longitudinal  groove.    Eyes  bare P.  catillifex 

22.  Upper  outer  orbits  and  vertex  separated  from  occiput  by  a  very 

fine  carina,  somewhat  interrupted  medially.   Body  8  to  11  mm. 

long P.  occidentalis 

Upper  outer  orbits  and  vertex  not  separated  from  occiput  by  a 

carina.    Body  7  to  8.5  mm.  long P.  bistriata 

Polybia  lugubris  H.  de  Saussure  (1854)  was  originally  described  as 
from  Guiana,  but  apparently  by  error.  The  species  has  only  been  taken 
definitely  in  southern  Brazil  and  is  unlikely  to  occur  in  the  Guianas. 
It  is  not  included  in  the  foregoing  key. 

Polybia  ujhelyii  Ducke  (1909)  is  known  from  the  Lower  Amazon 
(Brazil)  and  French  Guiana.    I  have  seen  it  from  Bolivia. 

Polybia  signata  Ducke  (1910)  is  known  from  French  Guiana,  Brazil 
and  Panama. 

Polybia  a  finis  R.  du  Buysson  (1908)  was  described  from  French 
Guiana.  Ducke  records  it  also  from  Brazil  and  Ecuador.  I  do  not 
know  this  species. 

Polybia  emaciata  Lucas  (1879).  This  is  in  my  opinion  the  oldest 
valid,  recognizable  name  for  the  well-known  South  American  wasp 
which  builds  a  nest  of  clay  with  a  circular  entrance.  It  is  generally 
called  Polybia  fasciata  H.  de  Saussure  (1854),  which  was,  however,  a 
misidentification  of  Polybia  fasciata  Lepeletier  (1836).  Lepeletier's 
fasciata  was  certainly  not  the  Polybia  with  the  clay  nest,  the  size  given 
being  much  too  large  (7  French  lines  =  15  mm.).  I  regard  P.  fasciata 
Lepeletier,  Vespa  fasciata  Olivier  (1791)  and  Vespa  fulvo-fasciata 
Degeer  (1773)  as  one  and  the  same  species  of  Stelopolybia.  Polybia 
caementaria  Ducke  (1904)  is  a  synonym  of  P.  emaciata.  The  species  is 
known  from  Dutch  Guiana,  French  Guiana,  Venezuela,  Colombia, 
Brazil,  Peru,  Bolivia,  Panama,  Costa  Rica,  Guatemala  and  the  Repub- 
lic of  Honduras. 


Polybia  gorytoides  Fox  (1898) 

British  Guiana:  west  bank  of  the  Demerara  River;  Bartica.  Also 
known  from  Brazil  (Oyapoc;  Amazon  Basin;  Chapada),  Dutch  Guiana 
(Kwakoegron,  Saramacca  River),  Bolivia  (Buena vista,  Dept.  Sta. 
Cruz)  and  Colombia  (Villavicencio,  Int.  del  Meta). 


282  bulletin:  museum  of  comparative  zoology 

Polybia  liliacea  (Fabricius,  1804) 

P.  liliacea  and  P.  striata  (Fabricius)  (Syn. :  P.  syncophanta  Gribodo) 
are  closely  allied  species,  with  the  same  type  of  coloration,  namely 
black,  with  extensive  pale  markings  on  thorax  and  abdomen  and  par- 
ticularly with  two  broad  longitudinal  pale  stripes  on  mesonotum. 
These  two  species  are  structurally  distinct  and,  as  Dr.  C.  Geijskes 
writes  me,  differ  in  the  nesting  habits. 

In  P.  liliacea  the  striation  and  puncturation  of  the  propodeum  are 
weak  or  obsolete,  the  concavity  showing  only  faint  traces  of  striae. 
The  postscutellum  is  rather  strongly  swollen.  The  first  abdominal 
tergite  is  short  and  much  swollen,  the  swelling  being  rather  abrupt  in 
profile,  the  basal  stalk-like  portion  well-defined  and  at  most  one-third 
of  the  total  length  of  the  tergite.  Usually  the  color  markings  are  very 
pale  yellow  or  ivory-yellow,  the  longitudinal  stripes  of  the  mesonotum 
often  coalescent  before  the  scutellum;  all  or  most  of  the  abdominal 
segments  bear  apical  bands.  This  type  of  coloration  was  correctly 
described  by  Fabricius  as  follows:  "Caput  cum  antennis  nigrum. 
Thorax  ater  limbo  lineolisque  duabus  crassis,  antice  abbreviatis, 
postice  coeuntibus,  scutello  lineolisque  duabus  sub  scutello  flavis. 
Abdomen  nigrum  segmentorum  marginibus  flavis.  Pedes  nigri." 

According  to  Dr.  Geijskes,  the  nest  of  P.  liliacea  is  very  long  (1.5 
m.)  and  sausage-shaped,  truncate  at  the  lower  end.  It  was  partly 
described  by  de  Saussure  (1858,  Et.  Fam.  Vesp.,  2,  pp.  cx-cxi)  and 
H.  Lucas  (1867). 

British  Guiana:  Warina;  Oronoque  River;  Georgetown;  Berbice 
Savannahs;  west  bank  of  Demerara  River;  Kartabo;  Bartica;  source 
of  Essequibo  River.  It  is  known  also  from  French  Guiana,  Dutch 
Guiana,  Brazil,  Venezuela,  Colombia,  Peru,  Bolivia,  and  Ecuador. 


Polybia  striata  (Fabricius,  1787) 

Polybia  sycophania  Gribodo  (1891)  I  regard  as  a  synonym  of 
Fabricius'  Vespa  striata.  Those  who,  like  Ducke,  refuse  to  recognize 
this  wasp  in  Fabricius'  description,  will  have  to  use  Gribodo's  name. 

In  P.  striata,  the  puncturation  of  the  propodeum  is  coarse  and  the 
striation  is  distinct  though  irregular  and  particularly  well  developed  in 
the  concavity.  The  postscutellum  is  moderately  swollen.  The  first 
abdominal  tergite  is  elongate  and  gradually  widened,  being  slightly 
swollen  in  profile,  with  the  basal  stalk  at  least  one-third  of  the  total 
length  and  poorly  defined.    Usually  the  color  markings  are  darker 


bequaert:  social  vespidae  of  the  gutanas  283 

yellow,  often  with  an  orange  tinge  and  the  stripes  of  the  mesonotum 
may  be  free  throughout.  The  apical  fascial'  of  the  abdomen  are  either 
as  in  liliacea,  or  much  reduced,  sometimes  almost  lacking.  Fabricius 
evidently  described  one  of  these  very  dark  specimens:  "Caput, 
abdomen,  pedes  nigra,  immaculata.  Thorax  niger  margine  antico 
tenuissime,  lineis  duabus  dorsalibus,  lateribus  baseos  obliquis,  scutello 
maculisque  sub  scutello  flavis.  Alae  albae  costa  fusca."  It  should  be 
noted  that  some  specimens  with  the  structural  characters  of  striata 
are  colored  exactly  like  liliacea,  so  that  the  color  is  not  a  reliable  char- 
acter.  On  the  other  hand,  I  have  seen  no  specimens  of  liliacea  either 
with  the  abdomen  almost  black  or  with  orange-yellow  markings. 

The  species  is  as  common  and  as  widely  distributed  as  P.  liliacea. 
I  have  seen  it  from  British  Guiana :  Kartabo  ;Warina.  Also  from  Dutch 
Guiana,  Trinidad,  Brazil,  Colombia,  Bolivia  and  Peru.  It  was  first 
described  from  French  Guiana. 

According  to  Dr.  Geijskes,  the  nest  of  P.  striata  is  a  white  spherical 
structure  hung  up  high  in  a  tree.  It  is  of  interest  that  the  Indians  in 
the  interior  of  Dutch  Guiana  distinguish  P.  striata  and  P.  liliacea  by 
name.   The  nest  of  P.  striata  has  not  yet  been  described. 

Ducke  (1910)  includes  P.  striata  and  P.  liliacea  in  his  key  among  the 
species  without  striation  on  the  propodeum.  As  the  striation  is  fairly 
distinct,  at  any  rate  in  P.  striata.  I  have  inserted  these  species  twice  in 
my  key. 

Polybia  jurixei  H.  de  Saussure  (1854) 

A  widely  distributed  species,  which  I  have  seen  from  British  Guiana : 
Kartabo.  It  is  also  known  from  French  Guiana,  Dutch  Guiana,  Brazil. 
Bolivia,  Peru,  Ecuador  and  Colombia  (Restrepo,  Int.  Meta) 

Polybia  bifasciata  H.  de  Saussure  (1854) 

This  species,  which  extends  from  Mexico  to  southern  Brazil,  is 
rather  variable  in  color.  In  a  recent  paper  (1943,  Jl.  New  York  Ent. 
Soc,  50,  for  1942,  pp.  300-303),  I  recognize  seven  forms  by  name. 
Only  two  of  these  have  been  taken  in  the  Guianas  thus  far.  Typical 
P.  bifasciata  I  have  seen  from  British  Guiana  (Kartabo;  Demerara 
River),  as  well  as  from  eastern  Peru.  The  var.  quadricincta  H.  de 
Saussure  (1854)  is  the  most  common  form  of  the  species.  It  occurs  in 
British  Guiana:  Bartica;  Kartabo;  source  of  Essequibo  River;  Warina, 
X.  W.  District.  I  have  also  seen  it  from  French  Guiana,  Trinidad, 
Venezuela,  Colombia,  Peru  and  Bolivia. 


284  bulletin:  museum  of  comparative  zoology 

Polybia  sericea  (Olivier  1791) 

Olivier's  typical  form  has  the  head  black,  the  thorax  and  first 
segment  of  the  abdomen  fulvous,  and  the  remainder  of  the  abdomen 
blackish.  This  is  a  common  wasp  throughout  most  of  Central  and 
South  America,  from  Guatemala  to  Buenos  Aires.  It  was  originally 
described  from  French  Guiana.  It  occurs  in  British  Guiana  (Upper 
Rupununi  River)  and  I  have  seen  it  from  Dutch  Guiana  (Paramaribo). 
Rhopalidia  rufithorax  Lepeletier  (1836),  Apoica  cubitalis  H.  de  Saussure 
(1854)  and  Polybia  melanocephala  Cameron  (1906)  were  based  upon 
typical  P.  sericea. 

Most  specimens  I  have  seen  from  British  Guiana  (Demerara;  Onver- 
wagt ;  Georgetown ;  Mt.  Roraima ;  Bartica  District)  belong  to  the  form 
of  the  species  with  both  first  and  second  abdominal  segments  fulvous. 
This  is  Polistes  nigripennis  Fabricius  (1804)  and,  if  a  name  is  needed  for 
it,  may  be  called  P.  sericea  var.  nigripennis  (Fabricius).  I  have  also 
seen  it  from  Venezuela. 

Polybia  chrysothorax  (Weber,  1801) 

Polistes  aurulenta  Fabricius  (1804)  and  Polybia  aurichalcea  H.  de 
Saussure  (1854)  are  synonyms. 

Common  in  British  Guiana :  Kartabo ;  Kalacoon ;  Demerara ;  Bartica 
District;  Georgetown;  Forest  Settlement,  Mazaruni  River.  I  have 
also  seen  it  from  Dutch  Guiana,  Colombia,  Brazil,  Venezuela  and 
Panama. 

Polybia  micans  Ducke  (1904) 

1.  Typical  P.  micans  is  russet  to  brownish-russet  with  more  testa- 
ceous areas,  particularly  at  the  apical  margins  of  the  abdominal  seg- 
ments and  the  base  of  the  second  tergite.  The  tegulae  and  legs  are 
russet-testaceous,  the  wings  strongly  tinged  with  russet.  In  British 
Guiana  it  was  taken  at  Kartabo  and  Bartica.  I  have  also  seen  it  from 
French  Guiana,  Dutch  Guiana,  Brazil,  Bolivia,  and  Ecuador.  It 
appears  to  be  somewhat  nocturnal  and  occasionally  flies  to  light. 
Polybia  sericeibalteata  Cameron  (1906)  I  regard  as  a  synonym  of 
typical  micans. 

2.  P.  micans  var.  velutina  Ducke,  was  described  by  Ducke  (1907) 
as  a  distinct  species,  but  I  am  unable  to  separate  it  by  structural  char- 
acters from  P.  micans.  It  is  fairly  uniformly  brownish-black,  the  tegu- 
lae and  legs  being  also  of  that  color.  There  are  a  few  testaceous  blotches 
on  the  head.   The  wings  are  more  yellowish  than  russet.   This  form  is 


bequaert:  social  vespidae  of  the  guianas  285 

rather  rare  in  British  Guiana:  Arakaka.   It  is  known  also  from  Brazil, 
Bolivia,  Peru,  Ecuador  and  Colombia. 

Polybia  dimidiata  (Olivier,  1791) 

British  Guiana:  West  bank  of  Demerara  River;  Oronoque  River, 
2°42';  Penal  Settlement,  Bartica  District.  I  have  also  seen  it  from 
Brazil,  Dutch  Guiana,  French  Guiana,  Bolivia,  Peru  and  Colombia. 

The  large  syrphid  fly,  Ceriodes  braueri  Williston,  is  perfectly  homeo- 
chromic  with  this  wasp. 

Polybia  rejecta  (Fabricius,  1798) 

I  recognize  three  color  forms  of  this  common  and  widely  distributed 
wasp,  but  transitional  specimens  are  frequent. 

1.  Typical  P.  rejecta  has  the  head,  thorax,  and  first  abdominal  seg- 
ment black,  with  very  few  pale  yellow  markings  (hind  margin  of 
postscutellum  and  narrow  apical  margin  of  first  tergite) ;  the  remainder 
of  the  abdomen  dull  brick-red,  with  or  without  narrow,  pale  margins 
(sometimes  ventrally  only).  Sometimes  the  red  turns  fuscous  or  black- 
ish. Common  in  British  Guiana:  Mt.  Everard;  Arakaka;  Amatuk; 
source  of  Essequibo  River;  Tumatumari,  Potaro  River;  west  bank  of 
Demerara  River.  Kartabo;  Bartica;  Rupununi;  Penal  Settlement, 
Bartica  District;  Moraballi  Creek,  Essequibo  River.  I  have  also  seen 
it  from  Dutch  Guiana,  French  Guiana,  Trinidad,  Venezuela,  Brazil 
and  Peru.  Polybia  bicolor  F.  Smith  (1857)  and  Eumenes  impunctus 
Provancher  (1888)  are  synonyms. 

Polybia  rejecta  race  javaryensis  Cameron  (1906)  is  a  species  of 
Mischocyttarus. 

2.  Var.  belizensis  Cameron  (1906).  Entire  body  black,  the  pale 
markings  of  the  thorax  as  in  the  typical  form;  abdomen  dorsally  with 
a  narrow  apical  margin  on  first  segment  only.  This  is  the  common 
form  in  Central  America,  where  I  have  seen  it  from  British  Honduras, 
Guatemala,  the  Republic  of  Honduras,  Costa  Rica,  and  Panama;  but 
it  occurs  also  in  Colombia,  Brazil  and  Peru. 

3.  Var.  litoralis  Zavattari  (1905).  Like  var.  belizensis,  but  all  or  most 
of  the  abdominal  tergites  with  rather  broad  pale  apical  margins. 
Described  from  Ecuador.  I  have  seen  it  from  Colombia,  where  it  is 
common;  but  Ducke's  locality  "Bogota"  is  due  to  an  erroneous  label, 
as  no  member  of  the  subfamily  Polybiinae  occurs  in  the  vicinity  of 
that  city. 


286  bulletin:  museum  of  comparative  zoology 

Polybia  occidentalis  (Olivier,  1791) 

The  most  common  social  wasp  of  tropical  America,  from  Mexico 
(northward  to  Nuevo  Leon  and  Tamaulipas)  to  northern  Argentina 
and  Uruguay.  In  the  Antilles  proper  it  is  known  only  from  St.  Vincent 
and  Grenada.  It  is  exceedingly  variable  in  size,  color  and  even  the 
shape  of  the  first  abdominal  segment,  but  all  of  the  forms  intergrade. 
The  following  seem  to  be  sufficiently  distinct  to  deserve  names. 

1.  Typical  P.  occidentalis  is  black,  with  few  or  many,  reduced  or 
more  extensive  pale  yellow  or  whitish  markings,  fairly  evenly  dis- 
tributed over  head,  thorax  and  abdomen;  most  or  all  of  the  abdominal 
tergites  have  pale  apical  margins;  mesonotum  usually  unstriped. 
Vespa  pygmaea  Fabricius  (1793),  Polybia  pygmaea  var.  minor  Moebius 
(1856),  P.  pygmaea  var.  major  Moebius  (1857),  P.  albo-picta  F.  Smith 
(1857),  P.  bohemani  Holmgren  (1808)  and  Eumenes  ductus  Provancher 
(1888)  were  based  on  some  of  the  variantswhich  I  include  undertypical 
occidentalis.  If  desired,  small  specimens,  with  more  slender  first  tergite 
and  few  white  markings,  might  be  called  var.  pygmaea  (Fabricius). 

This  form  extends  over  practically  the  entire  range  of  the  species 
and  is  common  in  the  Guianas.  British  Guiana:  Kartabo;  Bartica 
District;  Demerara;  Oko  River  (tributary  of  Cuyuni  River);  Upper 
Rupununi. 

Polybia  fastidiosuscula  var.  nigriceps  Zavattari  (1906)  appears  to  be 
based  upon  specimens  of  P.  occidentalis  much  like  the  typical  form,  but 
with  the  head  entirely  black  and  traces  of  pale  longitudinal  stripes  on 
the  mesonotum.  I  have  seen  such  specimens  occasionally  from  colonies 
of  otherwise  typical  occidentalis  and  cannot  regard  them  as  more  than 
individual  variants. 

Myraptera  elegans  Curtis  (1844)  was  somewhat  more  extensively 
bright  yellow,  particularly  on  the  head,  and  had  two  yellow  lines  on 
the  mesonotum.  It  is  doubtful  whether  it  should  be  given  varietal 
status.   It  is  clearly  transitional  to  var.  fiavifrons. 

2.  P.  occidentalis  var.  fiavifrons  F.  Smith  (1857)  has  the  head  almost 
entirely  yellow  (usually  with  an  orange  tinge)  and  is  also  extensively 
yellow  on  thorax  and  abdomen.  In  particular  the  mesonotum  bears 
two  broad  longitudinal  stripes,  often  fused  before  the  scutellum. 
Polybia  saussurei  Holmgren  (1808)  is  not  separable,  according  to  the 
types  I  saw  in  Stockholm.  This  form  is  homeochromic  with  the 
darker  specimens  of  P.  bistriata  (Fabricius)  (  =  occodoma  de  Saussure); 
but  the  structural  characters  given  in  the  key  readily  separate  the  two 
species. 


bequaert:  social  vespidae  of  the  guianas  287 

I  have  seen  the  var.  flavifrons  from  Ecuador  (Guayaquil),  Peru 
(Trujillo)  and  Brazil  (Coary,  Amazonas;  Lassance,  Minas  Geraes; 
Tres  Lagoas,  Matto  Grosso;  etc.). 

3.  Var.  parvula  (Fabricius,  1S()4i  was  originally  described  as  almost 
wholly  black,  with  only  a  narrow  pale  apical  margin  on  the  first 
abdominal  segment.  Many  transitions  connect  this  form  with  typical 
ocddentalis.  I  am  unable  to  separate  from  it  P.  occidentalis  var. 
diguetana  R.  du  Buysson  (1905). 

The  var.  parvula  often  occurs  in  the  same  districts  as  the  typical 
form,  but  is  relatively  rare  in  British  Guiana:  Demerara ;  Kartabo; 
Kaieteur,  Arakaka.  I  have  also  seen  it  from  Mexico,  Guatemala, 
Costa  Rica,  Panama,  Colombia,  Venezuela,  French  Guiana,  Dutch 
Guiana,  and  Brazil. 

4.  Var.  juruana  R.  v.  Ihering  (1904)  is  remarkable  for  the  extension 
of  the  yellow  markings  on  the  abdomen,  the  second  tergite  being  mostly 
of  that  color.  The  mesonotum,  however,  has  no  yellow  stripes.  I  have 
seen  it  from  Brazil  (Puerto  America,  Rio  Putumayo),  Bolivia,  Peru, 
and  Venezuela;  it  was  described  from  the  Jurua  River,  Brazil. 

Specimens  from  Aquidauana  (Matto  Grosso),  Brazil,  in  which  the 
apical  yellow  margin  of  tergite  2  is  considerably  widened  on  the  sides, 
but  narrow  in  the  middle,  connect  var.  juruana  with  typical  occident- 
alis. 

5.  Var.  scutellaris  (White,  1S41)  differs  from  typical  occidentalis  only 
in  the  yellow  being  restricted  to  scutellum  and  postscutellum,  which 
are  partly  or  wholly  of  that  color. 

I  have  seen  it  from  Brazil  (Bahia;  Sao  Paulo),  Paraguay  (Villarrica), 
northern  Argentina  (Entre  Rios)  and  Uruguay  (Montevideo). 

6.  Var.  spilonota  Cameron  (1904)  has  not  only  the  scutellum  and 
postscutellum  mostly  dark  yellow,  often  with  an  orange  tinge,  but  also 
a  large  orange-yellow  spot  (sometimes  divided)  on  the  propodeum,  as 
well  as  other  markings  on  head,  thorax  and  abdomen;  often,  but  not 
always,  the  mesonotum  bears  a  prescutellar  spot,  divided  anteriorly 
into  two  prongs;  the  markings  of  the  head  are  rather  small. 

It  appears  to  be  common  in  Nicaragua  (San  Marcos,  topo- 
types  taken  with  the  holotype,  but  probably  not  seen  by  Cameron; 
Managua;  Chinandega;  Granada).  I  have  also  seen  many  specimens 
from  Mexico,  Costa  Rica  and  Panama  which  provide  all  transitions 
between  typical  occidentalis  and  var.  spilonota,  sometimes  apparently 
in  the  same  nest. 

7.  Var.  ruficeps  Schrottky  (1902)  is  very  distinct,  the  head  being 
entirelv  or  mostlv  brick-red  to  dull  red,  with  or  without  vellow  mark- 


288  bulletin:  museum  of  comparative  zoology 

ings.  The  remainder  of  the  body  is  much  marked  with  yellow,  but 
without  mesonotal  stripes  and  with  the  apical  margins  of  the  tergites 
complete,  narrowed  or  interrupted  medially. 

It  is  common  in  northern  Argentina,  but  occurs  also  in  southern 
Brazil  (Corumba,  Matto  Grosso),  Paraguay,  and  Bolivia  (Buena 
Vista  de  Sta.  Cruz). 

Polybia  procellosa  Zavattari  (1906) 

Typical  procellosa,  known  from  Ecuador  only,  is  mostly  ferruginous- 
brown,  variegated  with  black  and  yellowish  markings,  the  legs  yellow- 
ish-russet, the  wings  gray  with  yellowish  tinge. 

The  var.  dubitata  Ducke  (1910)  is  black  with  a  few  whitish  markings, 
the  legs  black,  the  wings  grayish  without  yellowish  tinge.  It  was 
originally  described  from  the  Amazon  Basin  in  Brazil ;  but  I  have  seen 
it  from  British  Guiana  (Kurupung)  and  Peru. 

Polybia  bistriata  (Fabricius,  1804) 

The  species  generally  called  Polybia  oecodoma  H.  de  Saussure  (1854), 
is  Polistes  bistriata  Fabricius  (1804),  according  to  Fabricius'  types.  I 
regard  it  as  structurally  distinct,  not  as  a  color  variety  of  Polybia 
occidentalis.  The  characters  given  in  my  key  show  that  it  is  more 
closely  related  to  P.  catillifex.  Cameron's  Polybia  brunneiceps  (1912), 
described  from  British  Guiana,  without  more  definite  locality,  appears 
to  be  a  synonym  of  P.  bistriata.  The  ground  color  varies  from  blackish- 
brown  to  pale  russet. 

P.  bistriata  is  common  in  British  Guiana:  Kartabo;  Moraballi 
Creek,  Essequibo  River;  Rockstone,  Essequibo  River;  Demerara; 
Warina,  N.  W.  District;  Kalacoon;  Mt.  Roraima;  Penal  Settlement, 
Bartica  District.  I  have  also  seen  it  from  Brazil,  Dutch  Guiana, 
Venezuela,  Ecuador,  Peru,  and  Colombia. 

Polybia  catillifex  Moebius  (1856) 

British  Guiana:  Turesi  Falls;  Kamakusa;  Bartica;  Pakaraimo  Mts., 
head  of  Mazaruni  River.  Also  known  from  Dutch  Guiana,  Brazil, 
Ecuador,  Colombia,  and  Peru. 

Polybia  singularis  Ducke  (1909) 

British  Guiana:  Bartica  District;  source  of  Essequibo  River.  Known 
also  from  the  Amazon  Basin  in  Brazil,  Colombia  (Mapiri)  and  eastern 
Peru. 


bequaert:  social  vespidae  of  the  guianas  289 

As  shown  in  the  key,  this  is  closely  allied  to  P.  cmaciata.  It  builds  a 
similar  nest  of  clay,  but  with  a  long,  narrow,  slit-like  (not  a  circular) 
entrance.  Such  nests  were  described  by  F.  Smith  (1851)  and  H.  Lukas 
(1890),  but  without  knowledge  of  the  builder,  which  was  first  recog- 
nized by  Ducke  (1905;  erroneously  referred  to  P.  cmaciata). 

POLYBIA  TINCTIPENNIS  Fox  (1898) 

The  typical  color  form  of  this  species  is  known  from  southern  Brazil, 
eastern  Peru,  Honduras  and  Panama.  In  British  Guiana  (Kamakusa; 
Kartabo;  source  of  Essequibo  River)  it  is  replaced  by  the  var.  ncbulosa 
J.  Bequaert  (1943),  with  uniformly  infuscated  wings.  This  form  occurs 
in  Trinidad  and  probably  also  in  the  Amazon  Basin  of  Brazil. 

Polybia  rufitarsis  Ducke  (1904) 

I  have  seen  the  typical  form  of  this  species  from  British  Guiana: 
Moraballi  Creek,  Essequibo  River.   It  is  also  known  from  Brazil. 

Polybia  ignobilis  (Haliday,  1836) 

The  involved  synonymy  of  this  common  South  American  wasp  will 
be  discussed  elsewhere.  Polybia  atra  de  Saussure  (1854;  not  Vcspa 
atra  Olivier,  1791)  and  Polybia  nigra  de  Saussure  (1858)  are  synonyms. 

I  have  seen  a  few  specimens  from  British  Guiana:  Mt.  Roraima.  It 
is  known  definitely  also  from  Panama,  Colombia,  Venezuela,  Brazil, 
Paraguay,  Bolivia,  Peru  and  Argentina. 

Apoica  Lepeletier  (1836) 

There  is  only  one  structural  species  of  Apoica,  extremely  variable  in 
color. 

Apoica  pallida  (Olivier,  1791) 

A.  pallida  is  a  nocturnal  wasp,  frequently  attracted  by  light.  I  have 
recently  examined  several  thousands  of  these  wasps.  The  conclusions 
reached,  as  to  color  variation  and  distribution,  will  be  embodied  in  a 
paper  to  be  published  elsewhere.  I  am  able  to  distinguish  by  name  five 
color  forms,  all  of  which  occur  in  the  Guianas.  Although  each  form  is 
somewhat  variable,  there  are  few  truly  transitional  specimens.  The 
females  may  be  separated  by  means  of  the  following  key. 


290  bulletin:  museum  of  comparative  zoology 

1.  Body  testaceous,  pale  fulvous,  or  darker  brown,  either  uniformly 

so  or  with  part  to  most  of  the  abdomen  paler  than  head  and 
thorax.  No  or  very  few  pale  yellow  markings  (none  on  head  and 
mesonotum).   Wings  more  or  less  infuscated,  often  blackish.  .  .  . 

var.  thoracica 

Body  testaceous  to  pale  fulvous,  with  many  yellowish  markings  or 

extensively  whitish  or  yellow 2 

2.  Abdomen  (except  base  of  tergite  1)  dorsally  pale  yellow  or  nearly 

white,  often  with  silvery  pubescence  ("frosty");  venter  either 
yellow  or  pale  yellow  or  fulvous.  Head  and  thorax  pale  fulvous 
or  testaceous,  usually  spotted  with  yellow.    Wings  subhyaline, 

brownish-russet  along  the  costa var.  pallens 

Dorsum  of  abdomen  not  mostly  pale  yellow  nor  silvery 3 

3.  All  abdominal  tergites  with  a  pale  yellow,  narrow  or  wide,  apical 

margin;  sixth  tergite  entirely  yellow.  Head  and  thorax  spotted 
with  yellow  (mesonotum  often  with  two  or  four  yellow  stripes). 

Wings  subhyaline,  slightly  pale  russet  along  the  costa 

var.  arborea 
Abdominal  tergites  not  all  margined  with  pale  yellow 4 

4.  Abdomen  pale  fulvous  (except  first  tergite),  with  apex  of  first  tergite 

and  broad  base  of  second  tergite  pale  yellow.  Head  and  thorax 
blackish  without  yellow  spots  (not  stripes  on  mesonotum).  Wings 

strongly  infuscated,  nearly  violaceous-black var.  albi  macula 

Pale  fulvous  to  darker  brown.  Sometimes  with  apex  of  first  tergite 
spotted  with  yellow ;  sixth  tergite  as  a  rule  mostly  yellow.  Head 
and  thorax  with  many  pale  yellow  spots.  Mesonotum  with  longi- 
tudinal stripes  (at  least  traces).  Wings  slightly  russet,  darker 
along  costa typical  pallida 

1.  Typical  A.  pallida.  —  Syn.:  Vesta  pallida  Olivier,  1791;  Apoica 
lineolata  Lepeletier,  1836;  Polistcs  translucida  Spinola,  1851;  Apoica 
bilineolata  "Lepeletier"  H.  de  Saussure,  1854;  Apoica  lineata  "Lepele- 
tier" R.  du  Buysson,  1906. 

A  common  form  in  British  Guiana:  Rockstone,  Essequibo  River; 
Mackenzie,  Demerara  River;  Bartica;  Kartabo;  Upper  Rupununi 
River;  Demerara  River;  Tumatumari,  Potaro  River;  Torani  Ranch, 
Berbice  River;  Shudihar  River;  Wismar;  Penal  Settlement,  Mazaruni 
River;  Kamakusa.  Also  known  from  British  Honduras,  Trinidad, 
Dutch  Guiana,  Brazil,  Ecuador  and  Peru. 

2.  A.  pallida  var.  thoracica  R.  du  Buysson,  1906. 


N 


bequaert:  social  vespidae  of  the  guianas  291 

This  form  is  probably  as  common  as  typical  pallida.  British  Guiana : 
Mackenzie,  Demerara  River;  Tumatumari,  Potaro  River;  source  of 
Essequibo  River;  Kuyuwini  River;  Shudihar  River.  It  is  known  also 
from  Costa  Rica,  Panama,  Colombia,  Venezuela,  French  Guiana, 
Dutch  Guiana,  Brazil,  Bolivia  and  Peru. 

3.  A.  pallida  var.  pallens  (Fabricius).  —  Syn.:  Polistes  pallens 
Fabricius,  1804;  Apoica  pallida  Lepeletier,  1836. 

Very  common  in  British  Guiana:  Kamakusa;  Shudihar  River; 
Oronoque  River,  2°  42';  Kartabo;  Tumatumari,  Potaro  River;  Kuyu- 
wini River;  Georgetown ;  source  of  Essequibo  River.  I  have  seen  it  also 
from  British  Honduras,  Guatemala,  Nicaragua,  Costa  Rica,  Panama, 
Colombia,  Venezuela,  Trinidad,  Dutch  Guiana,  French  Guiana,  Brazil, 
Paraguay,  Bolivia,  Peru  and  Ecuador.  It  is  reported  from  southern 
Mexico. 

4.  A.  pallida  var.  arborca  H.  de  Saussure.  —  Syn. :  Apoica  arborea 
H.  de  Saussure,  1854. 

British  Guiana:  Kartabo;  Torani  Ranch,  Berbice  River;  Shudihar 
River;  Tumatumari,  Potaro  River;  Mazaruni  River.  Also  known  from 
Dutch  Guiana,  French  Guiana,  Brazil,  Bolivia,  eastern  Peru  and 
Ecuador. 

5.  A.  pallida  var.  albimacula  (Fabricius).  —  Syn.:  Polistes  albima- 
cula  Fabricius,  1804;  Polistes  albimaculata  "Fabricius"  H.  de  Saussure, 
1854. 

A  rare  form  known  only  from  Dutch  Guiana  and  British  Guiana: 
Shudihar  River;  Kartabo;  Pakaraimo  Mis.  at  the  headwaters  of  the 
Mazaruni  River. 

Stelopolybia  Ducke  (1910) 
(Including  Gymnopolybia  Ducke,  1914) 

Ducke  (1914,  Zool.  Lahrb.,  Abt.  Syst.,  36,  pp.  317  and  327)  divided 
his  genus  Stelopolybia  into  two  groups,  restricting  the  earlier  name  to 
the  species  which  build  nests  in  the  open,  but  with  the  combs  enclosed 
in  a  paper  envelope.  The  species  building  uncovered  combs  inside 
some  natural  cavity,  he  separated  as  Gymnopolybia.  Although  he 
mentioned  some  structural  differences  between  these  two  biological 
groups,  he  could  find  none  that  divided  the  species  either  consistently 
or  naturally. 

Polistes  angulata  Fabricius  (1804)  was  selected  as  the  type  of  Stelo- 
polybia by  R.  Lucas  (1912,  Arch.  f.  Naturgesch.,  77,  Bd.  4,  Heft  1, 


292  bulletin:  museum  of  comparative  zoology 

p.  210).1  Unfortunately  this  is  one  of  the  species  with  uncovered 
combs  which  Ducke  later  placed  in  Gymnopolybia.  O.  W.  Richards 
(1943,  Proc.  Ent.  Soc.  London,  ser.  B,  12,  pts.  3-4,  p.  45)  has  now 
chosen  Polybia  vulgaris  Ducke  (1904),  which  I  regard  as  a  synonym 
of  Vespa  fulvofasciata  Degeer  (1773),  as  the  type  of  Gymnopolybia. 
This  is  also  one  of  the  species  with  uncovered  combs.  It  thus  appears 
that  the  designated  genotypes  of  Stelopolybia  and  Gymnopolybia  are 
strictly  congeneric,  making  the  two  names  synonyms  and  leaving  the 
group  of  species  building  combs  inside  a  paper  envelope  without  a 
distinct  name.  I  feel,  however,  that  the  distinction  between  the  two 
groups  is  purely  biological.  The  morphological  differences  assigned  to 
them  by  Ducke  and  Richards  are  either  inadequate,  showing  transi- 
tions or  not  applying  to  all  the  species  of  each  group,  or  else  of  very 
secondary  importance  in  the  general  classification  of  the  Polybiinae. 
For  this  reason  I  do  not  propose  a  new  name  for  the  species  which  build 
combs  within  an  envelope. 

H.  de  Saussure  (1854)  divided  his  subgenus  Polybia,  s.  sir.  into  a 
number  of  divisions :  Alpha,  Iota,  Phi,  Mu,  Kappa  and  Omega.  Accord- 
ing to  F.  J.  Griffin  (1939,  Jl.  Soc.  Bibl.  Nat.  Hist.,  1,  pp.  211-212), 
these  were  all  published  the  same  year  (1854)  and  antedate  de  Saus- 
sure's  use  of  some  of  them  for  groups  of  solitary  wasps.  Only  Phi 
and  Mu  contained  species  now  placed  in  Stelopolybia,  in  addition  to 
others.  In  view  of  the  divergence  of  opinion  regarding  the  use  of 
Greek  letters  spelled  out  as  generic  or  subgeneric  names,  I  select  types 
which  will  make  them  synonyms  of  older  names.  Phi  originally  con- 
tained 10  species,  8  of  which  are  now  placed  in  Stelopolybia  and  one  in 
Mischoeyttarus,  one  being  unrecognized  though  probably  also  a  Stelo- 
polybia. I  herewith  designate  as  type  Vespa  phthisica  Fabricius  (1793), 
now  placed  in  Mischoeyttarus  subgenus  Kappa  de  Saussure  (1854)  (see 
O.  W.  Richards,  1941,  Proc.  Ent.  Soc.  London,  B,  10,  pp.  125-126). 
Mu  comprised  11  species,  6  now  placed  in  Polybia,  2  in  Stelopolybia 
and  3  in  Mischoeyttarus.  I  herewith  designate  as  type  Myrapetra 
scutellaris  White  (1841),  now  placed  in  Polybia  and  which  is,  more- 
over, the  monotype  of  Myrapetra  White  (1841). 

iR.  Lucas'  earlier  selection  of  P.  angulata  invalidates  O.  W.  Richards'  recent  (1943)  choice  of 
Polybia  infernalis  de  Saussure  as  the  type  of  Stelopolybia. 


bequaert:  social  vespidae  of  the  guianas  293 

Key  to  Guiana  Species 

1.  Dorsal  area  of  pronotum  rounded  off,  at  most  with  a  trace  of 

humeral  collar 2 

Pronotum  with  a  raised  humeral  collar,  sometimes  projecting  as 
lateral  angles.   Oculo-malar  space  long 5 

2.  Small  species,  10  to  12  mm.  long.   Eyes  with  rather  distinct  short 

hairs 3 

Large  species,  15  to  20  mm.  long.    Eyes  scarcely  hairy.    Oculo- 
malar  space  short,  but  distinct 4 

3.  Oculo-malar  space  very  long  (about  as  long  as  ninth  antennal  seg- 

ment).  Upper  plate  of  mesepisternum  longer  than  high 

S.  cajennensis 

Oculo-malar  space  very  short  (eye  almost  touching  mandibular 

condyle).   Upper  plate  of  mesepisternum  nearly  as  long  as  high 

S.  pollens 

4.  Clypeus  with  distinct,  fairly  large  punctures.    Anterior  margin  of 

pronotum  moderately  raised  near  the  coxae.  Lower  outer  orbit 
nearly  as  wide  as  eye  in  profile.   Larger,  wing  15  to  16  mm.  long 

S.  paracusis 

Clypeus  with  fine  punctures.  Anterior  margin  of  pronotum  strongly 

raised  into  a  translucent  lamella  near  the  coxae.    Lower  outer 

orbit  narrower  than  eye  in  profile.   Smaller,  wing  13  to  14  mm. 

long S.  obidensis 

5.  Sides  of  humeral  collar  strongly  projecting  and  angular.   Eyes  dis- 

tinctly hairy 6 

Sides  of  humeral  collar  rounded  off,  not  or  scarcely  projecting.  Eyes 
with  a  few  sparse  hairs 8 

6.  First  abdominal  segment  slender,  longer  than  the  combined  pro- 

podeum  and  postscutellum.    Wing  14  to  15  mm.  long,  slightly 

yellowish S.  constructrix 

First  abdominal  segment  scarcely  longer  than  the  propodeum ....  7 

7.  Body  orange-yellow  with  russet  areas,  the  hind  half  of  abdomen 

black;  second  tergite  with  yellow  base  and  apical  margin.   Wings 

strongly  russet-yellow,  17  to  18  mm.  long <S.  testacea 

Body  black  rarely  with  a  few  yellow  markings.   Wings  subhy aline, 
with  a  yellowish  tinge,  about  16  mm.  long S.  angulata 

8.  Larger;  wing  about  14  mm.  long.   First  abdominal  segment  rather 

abruptly  widened  and  bell-shaped,  slightly  angular  at  the  sides 
seen  from  above.  Second  tergite  rather  suddenly  widened  behind 
the  base S.  fuho-fasciata 


294  bulletin:  museum  of  comparative  zoology 

Smaller;  wing  9  to  11  mm.  long.  First  abdominal  segment  very 
gradually  widened,  not  bell-shaped.  Second  tergite  gradually 
behind  the  base S.  pallipes 

Stelopolybia  cajennensis  (Fabricius,  1798) 

Poly bia  lignicola  Ducke  (1904)  is  a  synonym. 

A  common  species  in  British  Guiana:  Bartica;  Kamakusa,  Warina, 
N.  W.  District;  Kartabo;  Forest  Settlement,  Mazaruni  River.  Also 
known  from  Dutch  Guiana  (Saint  Barbara  Plain,  Surinam  River; 
Paramaribo),  French  Guiana,  Trinidad,  Brazil,  Colombia,  Peru, 
Ecuador,  Panama,  and  the  Republic  of  Honduras. 

Stelopolybia  pallens  (Lepeletier,  1836) 

The  type  of  Rhopalidia  pallens  Lepeletier  is  now  in  Spinola's  collec- 
tion at  the  Turin  Museum,  where  it  was  recognized  by  Ducke  as  identi- 
cal with  Polybia  inf emails  de  Saussure  (1854).  Lepeletier 's  name  is 
valid;  he  described  his  R.  pallens  as  a  new  species,  without  any  refer- 
ence to  Polistes  pallens  Fabricius  (1804),  which  is  not  placed  now  in 
Stelopolybia.  Other  synonyms  are:  Polistes  rufina  "Illiger"  Erichson 
(1848),  Polybia  ampullaria  Moebius  (1856);  Eumenes  fiavopectus 
Provancher  (1888);  and  Polybia  internalis  Dalla  Torre  (1904). 

Common  in  British  Guiana:  Bartica;  Kamakusa;  Kaieteur;  Turesi 
Falls;  Kartabo;  Baracara,  Mazaruni  River;  Demerara;  Moraballi 
Creek,  Essequibo  River;  Courantyne  River ;  source  of  Essequibo  River; 
Forest  Settlement,  Mazaruni  River;  west  bank  of  Demerara  River. 
Also  known  from  French  Guiana,  Trinidad,  Brazil,  Bolivia,  Peru,  and 
Ecuador. 

Stelopolybia  paraensis  (Spinola,  1851) 

This  species  occurs  in  three  color  forms. 

1.  Typical  S.  paraensis  is  extensively  orange-yellow  or  pale  ferrugi- 
nous ventrally,  blackish-brown  or  black  dorsally  with  many  pale  yellow 
markings,  forming  longitudinal  stripes  on  the  mesonotum  and  apical 
fasciae  on  most  of  the  tergites;  base  of  first  and  second  tergites  also 
yellowish;  antennae  and  legs  orange-russet;  tegulae  yellowish;  wings 
strongly  tinged  with  yellow.  Not  rare  in  British  Guiana:  Mt.  Roraima 
Kalacoon;  Pakaraimo  Mts.,  head  of  Mazaruni  River;  Shudihar  River. 
Also  known  from  Brazil,  Peru,  and  Bolivia.  Homeochromic  with  S. 
obidensis. 


bequaert:  social  vespidae  of  the  guianas  295 

2.  S.  paraensis  var.  ruficornis  Ducke  (1905).  Mostly  black;  anten- 
nae russet  (scape  darker);  spots  on  sides  of  frons,  outer  orbits,  hind 
margin  of  pronotum,  tegulae,  longitudinal  streaks  on  propodeum, 
tibiae,  tarsi,  apical  margins  of  some  of  the  tergitcs  and  sternites,  more 
or  less  testaceous  or  whitish;  no  stripes  on  mesonotum.  Wings  as  in 
typical  form.  Originally  described  from  the  Upper  Amazon,  Brazil 
(Japura  River;  Tabatinga).  I  have  seen  it  from  Peru  (Iquitos;  Rio 
Tapiche).   Ducke  also  reports  it  from  eastern  Ecuador. 

Ducke  records  specimens  from  Chiriqui,  Panama,  transitional 
between  typical  paraensis  and  ruficornis.  I  have  seen  such  a  specimen 
from  Pozuzo,  Peru.  It  has  the  legs  mostly  yellowish  and  two  narrow 
yellowish  lines  on  the  mesonotum;  otherwise  it  is  like  ruficornis. 

3.  S.  paraensis  var.  (or  subsp.)  obscurior,  new. 

Female.  —  Body  black;  only  the  flagellum  (particularly  below)  and 
fore  tibiae  and  tarsi  somewhat  russet;  lower  inner  orbital  margins 
orange  or  dirty  yellow,  filling  the  ocular  sinuses.  Tegulae  black.  Wings 
as  in  typical  form. 

Ecuador:  Jatun  Yucu,  Rio  Napo  Watershed,  700  m.,  holotype  and 
paratypes  (W.  Clarke-Macintyre).  —  Colombia:  Restrepo,  Int.  Meta, 
500  m.,  paratype  (J.  Bequaert).  —  British  Guiaxa:  Mt.  Roraima, 
paratype  (J.  G.  Myers);  source  of  Essequibo  River,  paratypes  (J. 
Ogilvie).  —  Dutch  Guiana:  Brownsberg,  paratypes  (D.  C.  Geijskes). 
—  Holotype  and  paratypes  at  Mus.  Comp.  Zool.,  Cambridge,  Mass.; 
paratypes  at  U.  S.  Nat.  Mus.  and  Amer.  Mus.  Nat.  Hist. 

Perfectly  homeochromic  with  S.  angulata,  but  without  the  humeral 
angles  of  that  species. 


Stelopolybia  obidensis  (Ducke,  1904) 

Polybia  paraensis  var.  luctuosa  W.  A.  Schulz  (1905)  is  a  synonym. 

British  Guiana:  Tumatumari;  Pakaraimo  Mts.,  head  of  Mazaruni 
River;  source  of  Essequibo  River;  Kamakusa;  Tukeit;  Forest  Settle- 
ment, Mazaruni  River;  Turesi  Falls;  Kartabo;  Bartica;  Moraballi 
Creek,  Essequibo  River;  Penal  Settlement,  Bartica  District.  I  have 
seen  it  also  from  Dutch  Guiana  and  it  is  known  from  Brazil. 

Stelopolybia  constructrix  (de  Saussure,  1854) 

British  Guiana:  Pakaraimo  Mts.,  head  of  Mazaruni  River;  Kama- 
kusa ;  Kartabo ;  source  of  Essequibo  River.  Also  in  French  Guiana  and 
Brazil. 


296  bulletin:  museum  of  comparative  zoology 

Stelopolybia  testacea  (Fabricius,  1804) 

Common  in  British  Guiana:  Kartabo;  west  bank  of  Demerara 
River;  Mackenzie,  Demerara  River;  Bartica;  Pakaraimo  Mts.,  head 
of  Mazaruni  River;  Kuyuwini  River;  Oko  River,  a  tributary  of  the 
Cuyuni  River;  Mt.  Roraima.  Also  seen  from  French  Guiana,  Dutch 
Guiana,  Brazil,  Venezuela,  Bolivia  and  Peru. 

Stelopolybia  angulata  (Fabricius,  1804) 

The  typical  form  of  this  species,  without  yellow  markings  and  with 
black  tarsi,  occurs  in  British  Guiana:  Kaieteur;  Bartica;  Kamakusa; 
Arakaka;  west  bank  of  Demerara  River;  Warina,  N.  W.  District; 
Pakaraimo  Mts.,  head  of  Mazaruni  River;  Mt.  Roraima.  I  have  seen 
it  also  from  Venezuela,  Dutch  Guiana,  Brazil,  Ecuador,  Peru  and 
Bolivia.  In  Guatemala,  Costa  Rica,  Panama  and  parts  of  Colombia  it 
is  replaced  by  a  closely  allied  species,  S.  panamensis  (Cameron),  in 
which  the  humeral  angles  are  much  less  prominent. 

S.  angulata  var.  angulicollis  (Spinola,  1851)  has  the  tips  of  the 
femora  and  the  entire  tibiae  and  tarsi  yellowish ;  while  the  var.  ornata 
(Ducke,  1905)  has,  in  addition,  some  pale  yellow  spots  on  the  thorax. 
Neither  of  these  is  known  from  the  Guianas. 

Stelopolybia  fulvo-fasciata  (Degeer,  1773) 

I  regard  Polistes  hectica  Fabricius  (1804)  and  Polybia  vulgaris  Ducke 
(1904)  as  synonyms.  Vespa  ochrosticta  Weber  (1801)  may  also  have 
been  the  same.  Vespa  fasciata  Olivier  (1791)  appears  to  be  merely  a 
new  name  proposed  for  Vespa  fulvo-fasciata  Degeer.  Polybia  fasciata 
Lepeletier  (1836)  was  also  the  same  wasp. 

Common  in  British  Guiana:  Pakaraimo  Mts.,  head  of  Mazaruni 
River;  Kamakusa;  Warina,  N.  W.  District;  New  River,  a  tributary  of 
the  Courantyne;  Penal  Settlement,  Bartica  District;  source  of  Esse- 
quibo  River;  Rockstone,  Essequibo  River;  Mt.  Roraima;  Baracara, 
Mazaruni  River;  mouth  of  Meamo  River.  Also  known  from  French 
Guiana,  Dutch  Guiana,  Brazil,  Bolivia,  Peru,  Ecuador  and  Colombia. 

Stelopolybia  pallipes  (Olivier,  1791) 

The  unwarranted  emendation  "pallidipes"  was  first  used  by  Dalla 
Torre  (1894).  The  following  names  are  either  synonyms  or  based  upon 
color  forms:  Polybia  anceps  de  Saussure  (1854);  Polybia  lutea  Ducke 


bequaert:  social  vespidae  of  the  guianas  297 

(1904);  Polybia  myrmecophila  Ducke  (1905);  Polybia  festae  Zavattari 
(1906);  Polybia  pallipes  var.  centralis  Cameron  (1907);  and  Polybia 
pallipes  subsp.  cuzcoensis  Schrottky  (1911). 

1.  Olivier's  original  description  reads:  "Vespa  pallide  testacea, 
capite  thoracisque  dorso  nigro  maculatis,  ahdomine  fusco  apice  pallido. 
Elle  a  environ  cinq  lignes  de  long  [5  French  lines  =  12.3  mm.].  Les 
antennes  sont  noires,  avec  les  premiers  articles  d'un  fauve  pale  en 
dessous.  La  tete  est  d'un  fauve  pale,  avec  la  partie  superieure  tachee 
de  noir.  Le  corcelet  est  d'un  fauve  pale,  avec  trois  lignes  noires  sur  le 
dos.  Le  petiole  est  fauve  pale.  L'abdomen  est  obscur,  avec  la  base 
d'un  fauve  pale.  Les  pattes  sont  d'un  fauve  pale.  Les  ailes  sont  trans- 
parentes."  The  French  description  agrees  well  with  what  Ducke 
(1910)  called  " Stelopolybai  pallidipes"  and  de  Saussure  (1854,  Et. 
Fam.  Vesp.,  2,  PL  XXV,  fig  2)  figures  as  "Polybia  pallipes".  In  this 
typical  form  the  abdomen  has  the  first  tergite  (or  petiole)  pale  fulvous, 
the  second  yellowish  fulvous  with  infuscate  or  blackish  apical  margin, 
the  remaining  segments  blackish;  there  are  no  distinct  yellow  apical 
fasciae.    P.  lirfea  Ducke  does  not  seem  to  differ  from  typical  pallipes. 

2.  8.  pallipes  var.  anceps  (de  Saussure)  has  most  of  the  abdomen 
strongly  infuscate  or  blackish,  with  distinct  yellow  apical  margins  at 
least  on  tergites  2  and  3,  sometimes  also  on  4  and  5;  the  yellow  base  of 
the  second  tergite  is,  as  a  rule,  sharply  defined  from  the  fulvous  discal 
area. 

3.  S.  pallipes  var.  festae  (Zavattari)  is  the  extreme  melanistic  form 
of  the  species,  with  the  body  blackish-brown  or  black  and  few  yellow- 
ish markings  on  head  and  sides  of  thorax  (none  on  mesonotum  or 
abdomen).  It  is  known  only  from  Eastern  Ecuador  (San  Jose,  1800 
m.).   I  have  not  seen  it. 

4.  S.  pallipes  var.  cuzcoensis  (Schrottky)  has  the«black  abdomen  of 
var.  festae;  head  and  thorax  are  profusely  marked  with  yellow,  as  in 
the  typical  form.  It  appears  to  be  characteristic  of  parts  of  eastern 
Peru:  Achinamiza;  La  Chorrera,  Putumayo;  Chanchamayo.  I  have 
also  seen  it  from  Bolivia :  Huachi,  Rio  Beni.  It  is  evidently  the  form 
with  black  abdomen  mentioned  by  Ducke  from  the  Upper  Amazon: 
Tabatinga;  Iquitos. 

5.  S.  pallipes  var.  (or  subsp.)  fulvanceps,  new. 

Female.  Ground  color  orange  to  fuscous-yellow,  darker  on  the 
dorsum  of  the  abdomen,  more  yellowish  ventrally;  head  and  meso- 
notum with  the  same  blackish  markings  as  in  the  other  forms  of  the 
species;  narrow  base  of  second  tergite  and  broad  and  well-marked  apical 
margins  of  tergites  2  to  5  bright  yellow;  tergite  6  mostly  yellow. 


298  bulletin:  museum  of  comparative  zoology 

Wings  and  legs  as  usual.  This  form  combines  the  fulvous-orange 
abdomen  of  var.  myrmecophila  with  the  abdominal  bands  of  var. 
anccps. 

Colombia:  Rio  Frio,  Dept.  Magdalena,  holotype  and  paratypes 
(G.  Salt).  —  Peru:  Colonia  Perene,  paratype  (J.  C.  Bradley);  Puerto 
Bermudez,  Rio  Pichis,  paratypes  (J.  C.  Bradley);  Miriantiriani, 
Camino  del  Pichis,  paratypes  (J.  C.  Bradley).  —  Holot3rpe  and  para- 
types at  Mus.  Comp.  Zool.,  Cambridge,  Mass.;  paratypes  also  at 
Cornell  University  (Dept.  of  Entomology). 

6.  S.  pallipes  var.  myrmecophila  (Ducke)  has  the  head  and  thorax 
of  the  typical  form,  but  the  abdomen  is  almost  unicolorous  pale 
yellowish-russet  or  orange,  usually  with  more  yellow  bases  of  second 
and  third  tergites.  Sometimes  the  apical  margins  of  some  of  the  ter- 
gites  are  very  narrowly  or  faintly  yellowish.  P.  pallipes  var.  centralis 
Cameron  is  clearly  the  same  form,  as  was  recognized  by  Ducke. 

Typical  S.  pallipes  was  described  originally  from  French  Guiana 
(Cayenne)  and  I  have  seen  it  from  British  Guiana  (Mt.  Roraima), 
where  it  appears  to  be  uncommon.  I  also  know  it  from  Brazil,  Para- 
guay, and  Peru  (mouth  of  Rio  Cotuhe) 

The  var.  anceps  is  the  usual  form  in  British  Guiana:  Kamakusa; 
Kartabo;  west  bank  of  Demerara  River.  I  have  also  seen  it  from 
Dutch  Guiana,  Trinidad,  Brazil,  Venezuela,  Colombia,  Panama  and 
Bolivia. 

The  var.  myrmecophila  also  occurs  in  British  Guiana:  Source  of 
Essequibo  River;  Demerara  River;  Rupununi  River.  It  is  more  widely 
distributed  than  any  of  the  other  forms,  as  I  have  seen  it  from  Guate- 
mala, Honduras,  Costa  Rica,  Panama,  Colombia,  Ecuador,  Peru  and 
Brazil. 


Pseudopolybia  H.  de  Saussure  (1863) 

The  genus  was  revised  in  1938  (Rev.  de  Entomologia,  9,  pp.  112- 
113),  but  a  new  species  is  described  below.  Of  the  five  species  known 
at  present,  four  occur  in  the  Guianas. 

1.  Mesepisternum  not  divided  by  an  oblique  suture.  Humeral  margin 
of  pronotum  straight,  bluntly  ridged.  First  abdominal  segment 
longer  than  wide,  forming  a  broad  stalk.  Small;  fore  wing  5.5 
to  6  mm.  long P.  pusilla 

Mesepisternum  completely  divided  by  a  well-marked  oblique  suture 
into  an  upper  and  a  lower  plate 2 


bequaert:  social  vespidae  of  the  guianas  299 

2.  Small;  fore  wing  5.5  to  G.5  mm.  long.    First  abdominal  segment 

longer  than  wide,  forming  a  distinct  stalk,  gradually  widened 
from  base  to  apex  (seen  from  above).   Clypeus  slightly  produced 

medially P.  langi 

Larger;  fore  wing  11  to  12  mm.  long.  First  abdominal  segment 
either  cap-shaped,  or,  if  stalk-like,  not  gradually  widened  from 
base  to  apex.    Clypeus  strongly  produced  medially 3 

3.  First  abdominal  segment  slightly  longer  than  wide,  forming  a  broad, 

bell-shaped  stalk,  set  off  from  the  remainder  of  the  abdomen 

P.  difficilis 

First  abdominal  segment  wider  than  long,  cap^shaped  and  not  set 

off  as  a  stalk  from  the  remainder  of  the  abdomen ...  P.  vespiceps 

Pseudopolybia  vespiceps  (de  Saussure,  1863)  has  been  taken  in 
French  Guiana  and  Dutch  Guiana,  as  well  as  in  Brazil. 

Pseudopolybia  pusilla  (Ducke,  1904) 

British  Guiana:  Monkey  Jump,  Essequibo  River  (Oxford  Uni- 
versity Exped.). 

The  species  is  also  known  from  Brazil  (Para  and  Oyapoc).  It  is 
readily  mistaken  for  a  Leipomeles.  It  differs  from  L.  dorsata  in  the 
much  wider  clypeus  and  stronger  humeral  transverse  ridge.  The  nest 
is  as  yet  unknown. 

Pseudopolybia  difficilis  (Ducke,  1905) 

I  have  seen  this  species  from  British  Guiana:  source  of  Essequibo 
River.   It  is  also  known  from  Brazil,  Peru,  and  Bolivia. 

PsEUDOPOLYBtA  langi,  new  species.   Fig.  2 

Female.  Head  moderately  flattened,  slightly  wider  than  thorax; 
seen  in  front,  slightly  wider  than  high;  from  above,  rectangular  with 
receding  hind  corners,  about  twice  as  wide  as  long;  occipital  margin 
nearly  straight.  Vertex  and  genae  not  margined  by  a  carina  behind; 
gena  scarcely  narrower  than  the  eye  in  profile.  Oculo-malar  space 
about  half  the  length  of  the  fourth  antennal  segment.  Inner  orbits 
about  one  and  one-third  times  as  far  apart  on  vertex  as  at  clypeus. 
Ocelli  rather  large,  in  an  equilateral  triangle;  posterior  ocelli  about 
twice  as  far  from  eyes  as  from  each  other.  Interantennal  shield  broad, 
not  set  off,  strongly  but  bluntly  and  evenly  swollen.   Antennae  nearly 


300 


bulletin:  museum  of  comparative  zoology 


twice  as  far  apart  as  from  eyes.  Frons  slightly  convex.  Clypeus  at  its 
narrowest  about  one  and  two-thirds  times  as  wide  as  high,  irregularly 
pentagonal  with  the  upper  side  much  the  longest,  contiguous  to  the 
eyes  over  nearly  one-half  of  sides;  apical  margins  slightly  produced, 
ending  in  a  very  broad,  obtusely  rounded  point.  Mandible  with  an 
oblique  cutting  edge  of  four  sharp  teeth,  the  lower  three  subequal,  the 
upper  one  much  smaller;  most  of  outer  surface  flattened  or  slightly 
depressed.    Antenna:  scape  short  and  rather  thick,  scarcely  curved; 


Fig.  2.    Pseudopolybia  langi  J.  Bequaert.    Female.    A,  wings;  B,  head  in 
front  view;  C,  maxillary  palpus;  D,  labial  palpus;  E,  body  in  profile. 


second  segment  short,  moderately  swollen ;  third  less  than  three  times 
the  length  of  second;  fourth  and  fifth  about  as  long  as  wide;  sixth  to 
eleventh  slightly  wider  than  long;  twelfth  slightly  longer  than  wide 
at  base,  with  bluntly  rounded  tip;  flagellum  about  equally  thick 
throughout.  Maxillary  palpi  of  6  segments.  Labial  palpi  of  4  seg- 
ments, the  tip  of  the  third  with  a  short,  heavy,  erect,  curved  seta. 
Thorax  moderately  elongate;  in  profile,  about  one  and  one-third  times 
as  long  as  high ;  from  above,  nearly  twice  as  long  as  wide  before  tegu- 
lae.  Pronotum  evenly  rounded  into  a  semi-circle  anteriorly,  the 
humeral  margin  barely  indicated  by  a  very  weak  transverse,  blunt 


bequaert:  social  vespidae  of  the  guianas  301 

ridge  on  the  sides,  where  it  forms  no  angles ;  dorso-lateral  areas  sloping; 
lower  anterior  margin  of  ventrolateral  areas  with  a  vertical  swelling 
indented  by  a  deep  pronotal  fovea.  Mesopleuron  moderately  swollen ; 
mesepisternum  completely  divided  by  an  oblique  suture  into  an  upper 
and  a  lower  plate;  deep  mesepimeral  suture  ending  abruptly  and  far 
from  the  metapleuron.  Upper  sclerite  of  metapleuro'n  ending  obliquely 
below,  with  a  short  extension  along  mesometapleural  suture.  Scutel- 
lum  slightly  and  evenly  convex,  anteriorly  with  a  fine,  raised,  longi- 
tudinal line  which  continues  posteriorly  as  an  impressed  line.  Post- 
scutellum  transversely  elliptical,  moderately  swollen.  Propodeum  not 
swollen,  moderately  slanting  in  profile;  median  concavity  a  broad, 
shallow,  longitudinal  groove.  First  abdominal  tergite  narrowed, 
moderately  elongate,  very  gradually  widened  and  thickened  from  base 
to  apex,  where  it  is  less  than  half  as  wide  as  second  tergite,  triangular 
in  outline  from  above.  Second  tergite  from  above  wider  than  long, 
gradually  widened  at  base ;  in  profile,  slightly  and  evenly  convex.  Mid 
tibiae  with  two  spurs.  Claws  symmetrical,  unarmed.  Venation: 
second  cubital  cell  much  higher  than  wide;  third  cubital  longer  than 
wide,  rectangular,  about  as  long  on  cubitus  as  on  radius;  radial  cell 
very  long,  acute;  basal  vein  ending  in  subcosta  close  to  the  large 
stigma. 

Smooth  and  moderately  shiny,  without  appreciable  punctures  or 
other  sculpture.  Pubescence  sparse  and  short.  Eyes  with  many  dis- 
tinct erect  hairs. 

Head,  thorax  and  legs  pale  yellowish,  with  a  few  fuscous  areas  on 
frons,  vertex,  occiput,  pronotum,  mesepimeral  suture,  propodeum 
(concavity  and  extreme  sides),  hind  portion  of  postscutellum,  and 
tibiae.  Antennae  fuscous  above,  russet  below.  Teeth  of  mandibles 
fuscous.  Mesonotum  fuscous,  with  a  pair  of  broad  median  yellow 
stripes  and  a  shorter  yellow  streak  on  each  side  near  the  tegula. 
Abdomen  pale  fuscous  to  light  ferruginous,  more  or  less  yellowish  at 
the  base  of  the  second  tergite.  Wings  hyaline  throughout;  veins  pale 
fuscous;  stigma  translucent  medially. 

Length  (h.+th.+t.l+2):  5.5  mm.;  of  fore  wing,  5.6  mm. 

British  Guiana:  Kamakusa,  female  holotype  and  paratypes  from 
one  nest  (Herbert  Lang).  Holotype  and  paratypes  at  Mus.  Comp. 
Zool.,  Cambridge,  Mass.;  paratypes  also  at  Am.  Mus.  Nat.  Hist,  and 
U.  S.  Nat.  Mus. 

The  characteristic  thick  seta  of  the  labial  palpus  is  shorter  than  in 
some  other  species  of  the  genus  and,  being  of  a  pale  color,  is  more 
difficult  to  see. 


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Parachartergus  R.  v.  Ihering  (1904) 

This  genus  was  revised  by  me  in  1938  (Rev.  de  Entomologia,  9,  pp. 
105-112).  Unfortunately  I  called  it  there  Chartergus,  in  the  belief  that 
Ashmead  (1902)  was  the  first  to  select  a  genotype  for  that  name.  The 
late  Miss  Sandhouse  informed  me  that  Emile  Blanchard  in  1840 
(Hist.  Nat.  Ins.,  3,  Orth.  Nevr.  Hem.  Hym.  Lep.  Dipt.,  p.  395)  had 
designated  Vespa  nidulans  Fabricius  (1793),  a  synonym  of  Vespa 
chartaria  Olivier  (1791),  as  the  type  of  Chartergus  Lepeletier.  The 
name  Parachartergus  must  therefore  be  used  for  the  genus  which  I  had 
called  Chartergus. 

Key  to  Guiana  Species 

1.  Vertex  and  cheeks  not  separated  from  the  occiput  by  a  carina. 

Humeral  margin  of  pronotum  erect,  not  overlapping  nortouching 

the  vertex P.  frontalis 

Vertex  and  cheeks  separated  from  the  occiput  by  a  sharp  carina. 

Humeral  margin  of  pronotum  slanting,  touching  or  overlapping 

.  the  vertex 2 

2.  Vertex  on  each  side,  near  the  upper  inner  orbit,  with  a  slightly 

raised,  impunctate  area.   Smaller;  fore  wing  8  mm.  long 

P.  smithii 
Vertex  without  impunctate  lateral  areas 3 

3.  Body  thickset,  black.    Humeral  margin  of  pronotum  low,  ridge- 

like, opaque.  Wings  black,  with  the  apical  portion  either  some- 
what paler  or  decidedly  whitish.  Fore  wing  9  to  12  mm.  long.  .4 
Humeral  margin  of  pronotum  raised  into  a  high,  sometimes  trans- 
lucent lamella.  Wings  not  black  with  paler  or  whitish  tips. 
Fore  wing  6.5  to  8  mm.  long 5 

4.  Head  and  thorax  with  distinct,  long  erect  hairs P.  apicalis 

Head  and  thorax  covered  with  a  grayish  bloom,  the  erect  hairs 

restricted  to  clypeus,  postscutellum  and  propodeum 

P.  fratemus 

5.  Body  lengthened,  black  or  ferruginous.  Thorax  over  one  and  a  half 

times  as  long  as  wide  seen  from  above P.  fulgidipennis 

Body  short,  thickset,  mostly  testaceous  or  russet.    Thorax  about 

one  and  one-third  times  as  long  as  wide  seen  from  above 

P.  eolobopterus 

Parachartergus  eolobopterus   (Weber,   1801)   was  taken   in  French 
Guiana  and  is  known  also  from  Colombia,  Venezuela,  and  Trinidad. 


bequaert:  social  vespidae  of  the  guianas  303 

Parachartergus  frontalis  (Fabricius,  1S04),  not  yet  reported  from  the 
Guianas,  is  known  from  the  adjoining  Oyapoc  district  of  Brazil  and 
Venezuela. 

Parachartergus  apicalis  (Fabricius,  1804),  frequently  recorded  from 
the  Guianas,  possibly  does  not  occur  there,  as  it  is  often  confused  with 
P.fraternus,  which  I  regard  as  a  distinct  species. 

Paraciiartergus  fraterxus  (Gribodo,  1S91) 

1 .  The  typical  form  of  this  species  has  the  tips  of  the  wings  whitish 
with  pale  veins,  being  colored  exactly  like  P.  apicalis.  I  have  seen  it 
from  Dutch  Guiana  (Paramaribo). 

2.  In  the  var.  concolor  Gribodo  (1891),  the  tips  of  the  wings  are 
somewhat  paler  but  not  whitish  and  have  fuscous  veins.  I  have  seen 
it  from  British  Guiana  (Hepseba,  Courantyne  River;  Arakaka), 
French  Guiana,  Brazil,  Trinidad,  Colombia,  and  Panama.  It  was 
first  described  from  Venezuela. 

Parachartergus  fulgidipennis  (H.  de  Saussure,  1854) 

I  have  recognized  by  name  five  color  forms  of  this  species,  two  of 
which  occur  in  the  Guianas. 

1.  The  var.  griseus  Fox  (1898)  has  the  thorax  and  abdomen  black, 
the  abdomen  rather  dull  with  many  long  hairs,  and  the  head  partly 
pale  yellow.  The  wings  are  subhyaline,  broadly  black  along  the  costal 
margin,  without  distinct  cream-colored  transverse  patch.  Charter gus 
trichiosomus  Cameron  (1912),  described  from  British  Guiana,  is  a 
synonym.  I  have  seen  this  form  from  British  Guiana  (Moraballi 
Creek,  Essequibo  River),  Brazil,  and  Peru. 

2.  In  the  var.  fasdipennis  Ducke  (1905)  the  color  of  the  body  is  as 
in  var.  griseus,  but  the  abdomen  is  rather  shiny,  with  sparse  and  short 
hairs,  and  the  wings  have  a  distinct  cream-colored  transverse  patch, 
which  interrupts  the  black  streak  along  the  costal  margin.  It  was 
originally  described  from  Brazil;  but  I  have  seen  it  from  British 
Guiana:  Ite  Cattle  Trail. 

Parachartergus  smithii  (H.  de  Saussure,  1854) 

The  typical  form  of  this  species  was  taken  in  British  Guiana: 
Kartabo.  It  also  occurs  in  British  Honduras,  Costa  Rica,  Colombia, 
Ecuador,  Peru,  and  Brazil. 


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Leipomeles  Moebius  (1856) 
This  genus  is  monotypic. 

Leipomeles  dorsata  (Fabricius,  1804) 

W.  A.  Schulz  (1912,  Berlin.  Ent.  Zeitschr.,  LVII,  p.  87)  examined 
the  type  of  Polistes  dorsata  Fabricius  and  recognized  that  it  was  the 
wasp  described  by  Moebius  (1856)  as  Leipomeles  lamellaria.  Polybia 
nana  H.  de  Saussure  (1863)  and  Polybia  spilogastra  Cameron  (1912) 
are  other  synonyms. 

British  Guiana:  Source  of  Essequibo  River;  Monkey  Jump,  Esse- 
quibo  River.  Fabricius'  type  came  from  the  Essequibo  River.  Also 
known  from  Dutch  Guiana,  Brazil,  Bolivia,  Peru,  Ecuador,  and 
Panama  (Barro  Colorado). 


Bulletin  of  the  Museum  of  Comparative  Zoology 

AT    HARVARD    COLLEGE 

Vol.  XCIV,  No.  8 


THE  BIRD  FAUNA  OF  THE  WEST  SUMATRA  ISLANDS 


By  S.  Dillon  Ripley 

Division  of  Birds 
United  States  National  Museum 


With  Two  Plates 


CAMBRIDGE,  MASS.,  U.  S.  A. 
PRINTED    FOR    THE    MUSEUM 
October,;  1944  j 


PUBLICATIONS 
OF  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 
AT  HARVARD  COLLEGE 


The  Bulletin  and  Memoirs  are  devoted  to  the  publication  of 
investigations  by  the  Staff  of  the  Museum  or  of  reports  by  spec- 
ialists upon  the  Museum  collections  or  explorations. 

Of  the  Bulletin,  Vols.  I  to  XCIII,  and  Vol.  CXIV,  No.  1,  2, 
3,  4, 5,  6  and  7  have  appeared  and  of  the  Memoirs,  Vol.  I  to  LVI. 

These  publications  are  issued  in  numbers  at  irregular  intervals. 
Each  number  of  the  Bulletin  and  of  the  Memoirs  is  sold  separately. 
A  price  list  of  the  publications  of  the  Museum  will  be  sent  upon 
application  to  the  Director  of  the  Museum  of  Comparative 
Zoology,  Cambridge,  Massachusetts. 

After  1941  no  more  Memoirs  are  to  be  published. 


Bulletin  of  the  Museum  of  Comparative  Zoology 
AT    HARVARD    COLLEGE 

Vol.  XCIV,  No.  8 


THE  BIRD  FAUNA  OF  THE  WEST  SUMATRA  ISLANDS 


By  S.  Dillon  Ripley 

Division  of  Birds 
United  States  National  Museum 


With  Two  Plates 


CAMBRIDGE,  MASS.,  U.  S.  A. 

PRINTED    FOR    THE    MUSEUM 

October,  1944 


Xo.  8—  The  Bird  Fauna  of  the  West  Sumatra  Islands1 
By  S.  Dillox  Ripley 


CONTENTS 

Page 

Introduction 307 

Acknowledgements 308 

Ornithological  History 309 

Geography  of  the  islands 311 

A  Faunal  List  of  the  birds  of  the  islands 317 

Summary  and  Conclusions 415 

Bibliography 427 


INTRODUCTION 

The  islands  of  the  East  Indies  stretching  from  the  Asiatic  mainland 
down  to  the  southeast  for  more  than  two  thousand  m'les  'nto  the 
Papuan  and  Australian  regions,  form  a  true  paradise  for  the  student 
of  faunal  distribution  and  speciation.  The  position  of  this  myriad  of 
islands  large  and  small,  each  with  its  varying  type  and  degree  of  geo- 
graphic and  geologic  isolating  mechanisms  is  highly  interesting.  The 
equable  climate,  with  a  minimum  of  seasonal  or  spasmodic  changes, 
adds  to  the  effectiveness  of  even  the  most  limited  geographic  barriers. 
Ten  or  fifteen  miles  of  sea  is  enough  to  preclude  interchange  of  genes 
between  two  sedentary  populations.  As  an  example  of  this  kind  of 
speciation  the  bird  fauna  of  the  west  Sumatra  islands  offers  a  superla- 
tive opportunity  for  study. 

Several  factors  have  influenced  my  decision  to  study  the  birds  of  the 
west  Sumatra  islands.  During  1939  I  was  able  to  visit  Nias,  the  largest 
island  of  the  group,  where  I  made  a  small  collection  for  the  Academy 
of  Natural  Sciences  of  Philadelphia.  When,  early  in  1942  I  came  to  the 
United  States  National  Museum  from  Harvard,  I  found  that  I  would 
have  an  opportunity  in  Washington  to  make  an  exhaustive  study  of 
what  is  the  largest  and  finest  collection  of  birds  from  these  islands,  that 
of  the  late  Dr.  W.  L.  Abbott.  Dr.  Abbott's  collection  has  never  been 
reported  on  as  a  whole,  but  various  papers  by  the  late  Dr.  Charles 
Richmond,  Dr.  H.  C.  Oberholser,  and  the  late  Mr.  J.  H.  Riley  have 

1  In  the  absence  of  the  author  in  war  service,  the  proof  was  kindly  read  by  Dr.  Herbert 
Friedmann,  Curator  of  Birds,  U.  S.  National  Museum.     Editor. 


308  bulletin:  museum  of  comparative  zoology 

appeared  from  time  to  time  describing  more  than  one  hundred  and  thirty 
new  species  and  subspecies,  the  types  of  which  are  now  in  the  United 
States  National  Museum.  Thus  without  actually  returning  to  the 
islands,  now  out  of  the  question,  I  have  been  able  to  accumulate  re- 
markably complete  data  on  the  fauna  of  this  area. 


ACKNOWLEDGEMENTS  x 

To  the  late  Dr.  Glover  M.  Allen  of  Harvard,  my  councillor  and 
friend,  I  owe  a  great  deal  for  what  has  gone  into  this  study.  Professor 
A.  C.  Romer  has  been  most  helpful  with  aid  and  advice.  Mr.  James 
L.  Peters  has  been  kindness  itself  in  rendering  me  every  encouragement 
and  assistance.  Dr.  Ernst  Mayr  of  the  American  Museum  of  Natural 
History  and  Mr.  Rodolphe  de  Schauensee  of  the  Academy  of  Natural 
Sciences  have  freely  loaned  specimens  from  their  collections  and  have 
given  much  valuable  advice.  To  the  authorities  of  the  United  States 
National  Museum,  Dr.  Alexander  Wetmore,  Dr.  Herbert  Friedmann, 
and  Mr.  H.  G.  Deignan,  I  owe  a  tremendous  debt  of  gratitude  for  their 
generous  help  and  wise  counsel.  Certain  final  questions  of  identifica- 
tion remain  unsettled.  For  this  unfortunately  there  is  no  solution 
until  such  time  as  free  communication,  field  study,  and  research  in 
general  return  to  their  proper  places  in  this  now  sadly  maladjusted 
world.  That  this  will  happen  in  good  time  is  the  earnest  hope  and 
calm  conviction  of  all  those  of  us  who  now  labor  in  alien  and  exciting 
fields. 

S.  Dillon  Ripley 
Washington,  D.  C. 
November,  1942. 


>  Originally  presented  as  a  thesis  in  partial  fulfillment  of  the  requirements  for  the  degree  of 
Doctor  of  Philosophy  at  Harvard  University. 


RIPLEY:    BIRD    FAUNA    OF   THE    WEST    SUMATRA    ISLANDS  309 

ORNITHOLOGICAL  HISTORY 

The  first  serious  scientific  observing  and  collecting  on  the  west 
Sumatra  islands  was  done  by  Baron  H.  C.  B.  von  Rosenberg  who 
visited  Nias  in  1854.  So  far  as  is  known  the  list  of  56  species  of  birds 
published  by  Xieuwenhuisen  and  himself  in  1SG3  as  being  found  on 
Nias,  and  again  the  list  published  by  him  in  1878  of  60  species  of 
birds  noted  on  Nias  and  nearby  islands  are  not  based  on  actual  collected 
specimens  but  only  on  observations.  The  first  actual  collection  of 
birds  was  made  by  Signor  Elio  Modigliani  during  his  exploration  of 
Nias  from  April  through  August,  1886.  This  collection  of  62  species, 
eight  of  which  were  new,  was  published  on  by  Count  Salvadori  in  1886 
and  also  by  Modigliani  himself  in  1890.  By  this  collection  the  known 
avifauna  of  Nias  was  raised  to  102  species. 

Two  collections  were  made  among  these  islands  in  1S91.  One  on 
Nias  was  prepared  by  M.  J.  Claine  and  reported  on  by  Dr.  Oustalet 
in  1892.  Signor  Modigliani  meanwhile  had  gone  to  Enggano  and  his 
very  interesting  material  consisting  of  23  species,  seven  of  which  were 
new,  was  described  by  Salvadori  in  1892.  Another  collection  made  on 
Nias  was  that  of  Mr.  W.  Thomas,  a  missionary  on  the  island,  who 
sent  birds  to  Graf  von  Berlepsch.  Some  of  these  birds  were  described 
by  Salvadori  and  some  by  Berlepsch  in  1895.  Signor  Modigliani,  who 
was  still  in  the  islands,  now  visited  Sipora,  and  Salvadori  listed  his 
collection  from  there  of  34  species,  3  of  which  were  new,  in  1894. 

A  hard-working  and  able  young  entomologist,  Mr.  J.Z.Kannegieter, 
visited  Nias  and  the  Batu  Islands  during  the  winter  of  1895-96.  Mr. 
J.  R.  H.  Neervort  van  der  Poll,  for  whom  Kannegieter  was  working, 
presented  his  bird  collection  to  the  Leyden  Museum,  where  the  Nias 
collection  was  reported  on  by  Dr.  J.  Biittikofer.  Dr.  Biittikofer's  list 
raised  the  known  avifauna  of  Nias  to  128  species,  of  which  4  were 
described  as  new  in  his  paper  (1896).  Mr.  van  der  Poll  included  in- 
formation on  the  native  names  of  these  birds  as  well  as  notes  on  the 
color  of  the  soft  parts  of  the  specimens.  The  355  Batu  specimens, 
except  for  two  species  described  by  Finsch  (1899),  remained  unworked 
until  1940  when  the  Academy  of  Natural  Sciences  of  Philadelphia  pub- 
lished two  papers  by  de  Schauensee  and  Ripley,  and  de  Schauensee. 

The  next  collections  made  on  the  islands  were  those  of  the  tireless . 
Dr.  W.  L.  Abbott  who  voyaged  about  the  East  Indian  islands  for 
several  years  on  a  small  schooner  manned  by  an  adventuresome  and 
daring  Malay  crew.   Dr.  Abbott  started  in  the  autumn  of  1901  at  the 
northern  end  of  the  chain,  visiting  Simalur,  Babi,  Lasia,  and  the  islands 


310  bulletin:  museum  of  comparative  zoology 

of  the  Banyak  group,  Bangkaru  and  Tuangku.  A  year  later  in  the 
autumn  and  winter  of  1902,  Dr.  Abbott  along  with  Mr.  Charles  Boden 
Kloss  of  the  Federated  Malay  States  Museum  went  again  to  Simalur. 
From  there  they  sailed  south  to  the  Pagi  Islands,  the  Batu  Islands, 
Tello,  Tana  Massa,  and  Tana  Bala,  and  lastly  Nias.  Two  years  later 
in  the  fall  of  1904,  Dr.  Abbott  visited  Enggano.  His  last  trip  to  the 
group  was  to  Nias  in  the  spring  of  1905.  As  a  result  of  these  compre- 
hensive trips  more  than  1300  bird  skins  were  presented  to  the  United 
States  National  Museum. 

Several  papers  have  been  written  on  these  collections.  Dr.  Charles 
W.  Richmond  published  the  first  on  the  Simalur  trip  in  1903  describ- 
ing 19  new  forms.  Much  later,  Dr.  Harry  C.  Oberholser  (1919)  wrote 
up  Dr.  Abbott's  second  collection  on  Simalur  describing  one  new  form. 
In  these  two  papers  there  are  color  notes  on  the  species,  and  in  Dr. 
Richmond's  paper  there  are  some  short  field  notes  made  by  Dr.  Abbott. 
On  various  occasions  Dr.  Oberholser,  notably  in  his  well  known  paper, 
"Descriptions  of  One  Hundred  and  Four  New  Species  and  Subspecies 
of  Birds  from  the  Barussan  Islands  and  Sumatra"  (1912),  and  Mr. 
Riley  have  published  descriptions  of  new  forms  from  the  Abbott 
collection,  but  no  comprehensive  paper  on  the  collection  as  a  whole 
has  ever  appeared.  Mr.  Riley  at  the  time  of  his  death  was  working 
on  a  faunal  list  of  the  birds  of  these  islands.  It  is  unfortunate  that  he 
did  not  live  to  complete  it.  Needless  to  say  it  has  been  an  inestimable 
source  of  valuable  information  in  the  preparation  of  this  manuscript. 

The  next  collection  of  birds  from  these  islands  was  that  of  Dr.  E. 
Jacobson  and  Jonkheer  W.  C.  van  Heurn  made  in  1913  on  Simalur  and 
its  adjacent  islands.  Dr.  G.  C.  A,  Junge  published  on  this  collection 
in  1936,  describing  4  new  races  and  including  many  extremely  valuable 
field  notes  made  by  the  collectors.  There  is  also  a  good  deal  of  material 
about  the  eggs  of  some  of  these  forms,  which  were  collected  and 
measured. 

Due  to  difficulties  with  the  local  population,  Dr.  Abbott  was  not 
given  permission  by  the  Netherlands  authorities  to  visit  Siberut  and 
Sipora.  Later  in  1924  an  expedition  from  the  Raffles  Museum,  partly 
financed  by  funds  given  by  Dr.  Abbott,  visited  these  islands  and  a 
duplicate  set  of  these  birds  was  given  to  the  United  States  National 
Museum.  This  collection  was  reported  on  by  Messrs.  F.  N.  Chasen 
and  C.  Boden  Kloss  in  1926.  They  described  11  new  forms.  In  1927 
Mr.  Riley  described  3  more  forms  from  the  same  collection. 

So  far  as  can  be  discovered  the  next  ornithological  collecting  on  these 
islands  was  not  until  1937  when  three  collectors  were  in  the  field.  Miss 


Fig   1     Map  "i  Wat  Suraatni  l-lnrnf-,  -.|mumj»  depth  contours, 


RIPLEY:    BIRD    FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  311 

Barbara  Lawrence  made  a  collection  of  31  species  and  subspecies  from 
Nias  for  the  Museum  of  Comparative  Zoology  at  Cambridge,  Mass. 
Mr.  J.  J.  Menden  made  a  collection  on  North  and  South  Pagi,  part  of 
which  is  now  at  the  same  Museum.  The  third  collection  was  that  of 
Dr.  J.  K.  de  Jong  and  a  worker  from  Buitenzorg,  Saiin,  who  spent  two 
months  on  Enggano.  The  first  two  collections  have  not  been  worked 
up,  but  the  Enggano  birds  were  reported  on  by  Dr.  Junge  (1938),  add- 
ing 5  species  to  the  list  of  birds  found  on  that  island.  In  June  1939  I 
spent  a  few  days  on  Nias  and  secured  specimens  representing  45  species 
and  subspecies.  This  material  has  been  written  up  by  de  Schauensee 
and  myself  (1940),  and  four  new  forms  were  described. 

Altogether  some  two  hundred  twenty  species  and  subspecies  have 
been  described  from  the  islands  of  which  the  present  paper  recognizes 
one  hundred  fourteen  forms.  The  total  list  of  the  bird  fauna  of  this 
small  archipelago  runs  to  two  hundred  eighty  species  and  subspecies 
including  migrants.  With  the  exception  of  three  races  I  have  been 
able  to  see  specimens  of  all  the  endemic  forms  occurring  in  this  group. 
These  three  races  are:  Spizaetus  cirrhotics  vanheurni  and  Eurostopodus 
macrotis  jacobsoni  from  Simalur,  and  Coraeina  striata  kannegieteri 
from  Nias.  Present  conditions  indicate  that  collecting  will  be  impos- 
sible on  these  islands  for  some  time.  However,  I  think  that  except  for 
the  recording  of  more  migrants  and  the  extension  of  range  of  some 
known  forms  the  present  list  should  stand  for  some  time  to  come.  I 
do  hope  that  in  the  future  there  will  be  an  opportunity  for  more 
biological  field  work  to  be  done  on  the  birds  of  these  islands.  Un- 
doubtedly they  serve  as  a  splendid  natural  speciation  laboratory. 

GEOGRAPHY  OF  THE  ISLANDS 

The  west  Sumatra  Islands  lie  in  a  chain  for  a  distance  of  approxi- 
mately six  hundred  miles  paralleling  the  western  coast  of  the  island  of 
Sumatra.  This  chain  runs  roughly  from  northwest  to  southeast  ex- 
tending from  Lat.  3°N„  Long.  95°35'E.,  to  Lat.  5°30'S.,  Long. 
102°24'E.  The  islands  have  never  been  carefully  explored  by  a 
geologist.     (See  map,  fig.  1). 

The  original  appearance  of  these  islands  is  presumably  due  to  the 
same  phenomena  which  caused  the  rise  of  the  Barussan  Mountains  in 
western  Sumatra.  Brouwer  (The  Geology  of  the  Netherlands  East 
Indies,  1925,  p.  2)  has  this  to  remark  about  these  mountains: 

"Sumatra  lies  along  the  axis  of  a  great  geanticline.  Stretching  along 
the  western  part  of  the  island,  near  to  the  coast  are  the  Barissan  [sic] 


312  bulletin:  museum  of  comparative  zoology 

Mountains  which  give  to  this  side  of  the  island  a  rough  and  rugged 
character.  .  .  .  The  present  Barissan  Mountains  are  associated  with 
late  Tertiary  and  post-Tertiary  mountain-building.  Considerable  vul- 
canism  occurred  during  a  great  part  of  the  Tertiary  period  and  even 
yet  there  are  many  active  volcanoes."  Later  (p.  84),  he  says:  "The  row 
of  islands  to  the  west  of  Sumatra  is  known  for  its  frequent  and  intense 
earthquakes,  but  volcanoes  are  entirely  lacking." 

Professor  G.  A.  F.  Molengraaff  in  an  interesting  article  ("Modern 
Deep-Sea  Research  in  the  East  Indian  Archipelago",  Geog.  Journ. 
57,  No.  2,  Feb.  1921,  p.  95.)  has  a  good  deal  to  say  about  the  forma- 
tion of  the  west  Sumatra  Islands.  His  discussion  of  folding  (p.  108) 
follows : 

"As  soon  as  the  upper  portions  of  the  folds  which  develop  at  a  cer- 
tain depth  approach  the  Earth's  surface  and  the  majority  of  the  rocks 
under  diminished  pressure  can  no  more  be  folded  without  being  frac- 
tured (van  Hise's  zone  of  fracture),  the  continuity  of  the  strata  will  be 
broken  and  the  culminating  portions  of  the  elevating  anticlinal  axes 
will  be  fractured  and  show  at  the  surface  as  isolated  portions  or  blocks, 
their  extent  and  shape  being  greatly  dependent  on  the  geological 
structure  and  the  differences  in  rigidity  of  the  composing  rocks.  This 
may  suffice  to  explain  why  an  elevating  submarine  ridge  formed  by 
an  anticlinal  axis  will  appear  at  the  surface  as  a  row  of  blocks,  i.e. 
islands  separated  by  deep  channels. 

"In  the  great  geosynclinal  area  between  the  continents  of  Asia  and 
Australia  one  arc  of  folding  belonging  to  the  Alpine  system,  and  known 
as  the  Malay  arc,  appears  to  originate  from  the  Burma  arc,  .  .  .  and 
can  be  followed  .  .  .  frcm  the  extreme  north-western  end  of  Sumatra, 
through  this  island  and  the  island  of  Java.  .  .  as  far  as  the  Banda  sea. 

"In  its  western  section  where  it  borders  on  the  Indian  Ocean,  as  in 
the  central  portion  of  the  island  of  Sumatra,  this  arc  now  consists  of 
two  major  folds,  as  is  illustrated  by  an  ideal  cross  section  (Fig.  2) 
from  the  Indian  Ocean  towards  the  stable  portion  of  the  Sunda  land. 
It  would  show  the  following  sequence: 

1.  Indian  Ocean. 

2.  Sunda  trough,  first  geosyncline. 

3.  Range  of  coastal  islands  girdling  the  west  coast  of  Sumatra  in- 
cluding the  Mentawei  islands,  first  geoanticline. 

-  4.  Mentawei  trough  and  corresponding  trough-shaped  depths, 
second  geosyncline. 

5.  Non-volcanic  and  volcanic  mountain  ranges  of  Sumatra  and 
Java,  second  geoanticline. 


RIPLEY:    BIRD    FAUNA    OF   THE    WEST    SUMATRA    ISLANDS  313 

6.  Tertiary  terrain,  folded  in  late  Tertiary  and  early  Pleistocene 
time,  now  practically  stable,  third  geosyncline. 

7.  Stable  Sunda  land  including  the  Sunda  shelf." 


wsw. />/  ^^  ^  *£A  LCVEL ENE 


1.         2.  3.4.         5.  <o.  7. 

Fig.  2.  Schematic  representation  of  late  Tertiary,  Pleistocene,  and  post- 
Pleistocene  crustal  movements  in  the  western  section  of  the  Malay  geosynclinal 
area.    (After  Molengraaff)   The  numbers  refer  to  those  listed  above. 

The  northern  group  of  islands  consists  of  Simalur  and  its  outliers. 
Simalur  is  about  fifty-five  miles  long,  varying  in  width  from  five  to 
fourteen  miles.  It  lies  about  sixty-five  miles  from  the  coast  of  Sumatra. 
It  is  a  rocky  well-wooded  island,  the  main  mountain  of  which,  Sibalur, 
rises  to  625  metres  in  height.  Simalur  is  thinly  populated  and  there  are 
few  gardens  or  clearings.  Besides  raising  sago,  coconuts  and  vegetables 
for  their  own  subsistance,  the  main  industry  of  the  people  is  raising 
water  buffalo.  The  principal  village  is  Sinabang  in  the  southeastern 
part  of  the  island. 

Simalur  is  outside  the  two  hundred  metre  line  which  curves  out  from 
the  Sumatran  coastline  and  envelops  the  Banyak  islands  to  the  south- 
east. Simalur  lies  on  its  own  two  hundred  meter  bank  which  encloses 
several  nearby  islands.  Northwest  lie  the  Kokos  or  Sa  Laut  Islands, 
two  small  flat  islets  which  are  the  most  northerly  extensions  of  the 
entire  archipelago.  These  two  islets  are  two  miles  by  one  and  a  half 
miles,  and  one  thousand  yards  in  diameter  respectively.  They  are 
connected  with  Simalur  twenty-five  miles  to  the  south  east  by  a  sub- 
terranean bank  less  than  thirty-six  metres  in  depth.  They  are  mainly 
covered  with  coconut  trees  and  the  only  collectors  to  visit  them  were 
Jacobson  and  van  Heurn  who  spent  a  day  there,  August  21,  1913. 
They  found  two  forms  on  Kokos,  Caloenas  nicobarica  nicobarica  and 
Eudynamis  scolopacea  simalurensis.  Presumably  other  birds,  particu- 
larly migrant  shore  birds,  occur  there  from  time  to  time. 

The  only  other  islets  which  have  been  collected  on  in  the  immediate 
vicinity  of  Simalur  are  Djawi  Djawi,  Lugu,  and  Pulu  Pandjang  in 
Sinabang  harbor,  and  Siumat,  six  miles  off  shore  in  an  easterly  di- 
rection. Siumat  is  about  two  and  a  half  miles  long  with  some  original 
forest  as  well  as  coconut  palms.  Both  Dr.  Abbott  and  Jacobson  and 
van  Heurn  collected  there  getting  three  common  species  of  small  island 


314  bulletin:  museum  of  comparative  zoology 

birds:  Sterna  dougalli  bangsi,  Chalcophaps indica  indica,  and  Halixetus 
leucogaster. 

South  of  Simalur  by  about  fourteen  miles  are  two  small  islands 
separated  from  each  other  by  a  strait  one  and  one  quarter  miles  wide. 
They  are  sometimes  called  the  Tapah  islands.  Both  are  small  low  coral 
islands  covered  with  original  forest  and  uninhabited.  Lasia,  the  north- 
ernmost is  smaller,  less  than  two  miles  in  length,  and  surrounded  by 
fringing  reefs.  Babi,  the  larger,  is  about  seven  miles  in  diameter  and 
nearly  round.  Both  islands  lie  on  a  shelf  separated  from  the  neighbor- 
ing islands  to  the  north,  east,  and  south  by  depths  greater  than  two 
hundred  meters.  The  only  collector  to  visit  Babi  and  Lasia  has  been 
Dr.  Abbott.  Together  these  two  small  islands  have  three  endemic 
races  not  found  on  any  of  the  other  islands  in  the  group.  These  are : 
Psittacula  alexandri  major,  Coracina  striata  babiensis,  and  Hypothymis 
azurea  abbotti. 

The  Banyak  Islands  lie  about  thirty-eight  miles  off  the  coast  of 
Sumatra,  and  forty-five  miles  south-east  of  Simalur.  Bangkaru,  the 
most  westerly,  is  thirty  miles  east  of  Babi.  The  principal  islands  of 
the  more  than  fifty  islets  and  reefs  in  the  group  are :  Tuangku,  seven- 
teen miles  long  by  five  miles  wide,  Bangkaru  with  an  area  of  twenty 
square  miles,  and  Ujung  Batu,  a  small  narrow  island.  These  islands 
are  decidedly  hilly  and  well  forested.  A  mountain  on  Bangkaru, 
Amintolan,  reaches  a  height  of  three  hundred  three  meters.  There  is 
a  hill  at  the  south  end  of  the  island.  In  between  the  land  is  very  low 
so  that  at  any  distance  the  appearance  presented  is  that  of  two  islands. 
Tuangku  has  a  mountain  of  three  hundred  thirteen  meters  on  the  north 
coast.  The  eastern  coast  is  low  and  covered  with  mangroves.  There  is 
a  small  population  on  these  islands  of  which  more  than  half  (536  in 
1911)  live  in  Tuangku. 

The  only  collections  from  the  Banyak  islands  have  been  those  of 
Dr.  Abbott  who  visited  Tuangku  and  Bangkaru.  Richmond  (1903, 
p.  485)  notes  that  more  species  were  seen  on  Tuangku  than  Dr.  Abbott 
had  found  on  Simalur.  However,  Abbott  noted  that  "no  large  parrots, 
hornbills,  or  barbets  were  seen  or  heard,  and  no  drongos  or  orioles 
were  noticed." 

The  largest  island  of  the  group  is  Nias  which  is  seventy  miles  long 
and  from  twelve  to  twenty-two  broad.  It  lies  twenty-eight  miles  south 
of  the  Banyak  islands  and  about  fifty-six  miles  south  west  of  Ujung 
Singkel,  a  projecting  point  of  the  Sumatra  coastline.  Nias  lies  on  its 
own  two  hundred  meter  shelf.  The  nearest  point  of  contact  is  with 
that  part  of  the  Sumatran  shelf  which  extends  south  of  the  Banyak 


RIPLEY:   BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  315 

islands.  The  elevation  of  Nias  reaches  a  height  of  eight,  hundred  eighty- 
six  meters.  In  general  the  central  part  of  the  island  is  rather  high 
forming  an  irregular  plateau  which  extends  down  on  all  sides  gradually 
to  the  coast.  The  general  impression  is  of  a  drowned  island  with  an 
irregular  sloping  outline  surrounded  by  outlying  submerged  rocks  and 
coral  reefs  in  all  directions.  Further  out  from  the  shore  are  many 
small  islands  such  as  the  Hinako  group  which  lie  on  a  ten  fathom  bank 
extending  out  from  the  west  coast  of  Nias.  Presumably  most  of  these 
islands  have  been  connected  with  Nias  at  one  time  or  another. 

Nias  has  the  largest  population  of  any  of  the  islands  numbering 
about  one  hundred  thousand  at  the  last  census.  The  main  centers 
are  in  the  southern  part  of  the  island  where  there  are  many  large 
villages.  This  southern  part  of  the  island  has  been  rather  heavily 
deforested  with  the  result  that  original  forest  is  now  found  only  in  a  few 
places  on  the  hill  tops.  The  northern  part  of  the  island  on  the  other 
hand  is  populated  by  a  somewhat  less  aggressive  people  whose  culture 
apparently  does  not  demand  the  construction  of  the  giant  houses  found 
to  the  south. 

In  this  part  of  the  island  much  original  forest  remains  and  there  are 
presumably  several  new  records  to  be  added  to  the  bird  list  of  the  island 
as  many  fewer  collectors  have  worked  here.  Nias  has  been  by  far  the 
best  collected  of  all  the  islands  due  to  its  size  and  accessibility.  Until 
1942  it  was  the  only  one  of  the  islands  which  was  visited  regularly  by 
an  inter-island  steamer  sailing  from  Sibolga  to  Goenong  Sitoli  every 
two  weeks.  The  main  trade  of  the  island  was  in  pigs  which  were  much 
in  demand  in  the  Chinese-populated  centers  on  Sumatra. 

Southeast  of  Nias  by  about  forty-five  miles  lie  the  Batu  Islands. 
These  consist  of  three  large  islands,  Pini,  Tana  Massa  and  Tana  Bala 
surrounded  by  more  than  twenty  smaller  ones.  Besides  these  three 
large  islands,  Tello  and  Lago  are  the  only  islands  that  have  been  visited 
by  collectors.  Tana  Massa  is  twenty -seven  miles  long  by  five  miles 
broad.  Tana  Bala  is  twenty-two  miles  long  by  seven  broad.  Pini  is 
twenty  miles  long  and  six  broad.  All  these  islands  are  rather  low  and 
without  distinguishing  features.  The  population  is  small  and  the  origi- 
nal forest  largely  remains.  Tello,  less  than  two  miles  long,  is  the  seat 
of  administration  and  the  most  populated  island  in  the  group.  Lago  is 
a  small  coconut  covered  islet  three  miles  northeast  of  Tana  Massa. 
Most  of  these  islands  are  surrounded  by  reefs  sometimes  five  or  six 
miles  out  in  all  directions.  The  whole  group  lies  within  the  two  hundred 
meter  line  which  curves  out  from  the  Sumatra  mainland  and  continues 
southeast  in  a  narrow  belt  inclosing  the  islands  below  the  Batu  group. 


316  bulletin:  museum  of  comparative  zoology 

The  name  for  those  islands  to  the  south  of  Tana  Bala  is  Mentawi. 
This  includes  Siberut,  Sipora,  North  and  South  Pagi,  and  the  adjacent 
small  islands.  North  and  South  Pagi  by  themselves  are  called  the 
Pagi  or  Pagai  islands.  All  four  of  these  islands  are  inhabited,  hilly, 
and  evidently  of  volcanic  formation.  Although  these  islands  lie  on  a 
relatively  shallow  bank  connected  with  Sumatra  to  the  north,  there  is 
a  definite  gap  directly  to  the  east  towards  the  nearest  part  of  the  Su- 
matran  mainland.  Here  there  is  the  deep  Mentawi  basin  reaching  a 
depth  of  over  sixteen  hundred  meters.  Thus  except  for  the  one  link  to 
the  north  there  is  no  evidence  to  indicate  that  the  Mentawi  group  has 
ever  been  connected  with  Sumatra. 

Siberut,  thirty  miles  to  the  south  of  Tana  Bala,  is  sixty  miles  long  by 
fifteen  to  twenty-four  miles  in  breadth.  It  is  heavily  wooded  with  ex- 
tensive marshes  along  the  coast.  There  are  about  eight  thousand  in- 
habitants at  an  extremely  primitive  cultural  level.  The  island  is  of  all 
the  group  perhaps  the  least  known  and  least  explored.  The  only 
collecting  of  birds  and  mammals  has  been  that  of  Messrs.  Kloss  and 
Smedley  in  1924. 

Sipora  is  a  small  densely  overgrown  island  about  thirty-three  miles 
long.  The  low-lying  land  is  extremely  marshy,  so  much  so  that  in 
former  years  the  best  communication  into  the  interior  was  by  canoe. 
The  highest  point  on  the  island  is  three  hundred  thirteen  meters. 
Sioban,  the  main  village  is  on  the  eastern  shore  and  is  a  small  copra 
shipping  port. 

Both  North  and  South  Pagi  present  a  slightly  different  picture  from 
Siberut  and  Sipora.  The  Pagi  islands  rise  more  steeply  from  the  sea, 
so  much  so  that  the  ten  fathom  curve  runs  very  close  along  the  shore. 
There  are  a  few  villages  principally  along  the  east  coast  but  the  popu- 
lation is  not  large.  Apparently  the  islands  are  principally  covered  with 
original  forest.  North  Pagi  lies  about  twelve  miles  south  of  Sipora. 
There  are  very  few  inhabitants  and  little  is  known  of  the  islands. 
Except  for  the  collection  made  by  Dr.  Abbott  and  Mr.  Kloss  in  1902 
there  has  been  only  the  short  trip  of  Mr.  Menden  in  1937.  It  is  unfor- 
tunate that  no  field  notes  were  made  on  these  two  visits,  for  undoubt- 
edly the  Pagi  Islands  are  extremely  interesting  from  the  faunistic  and 
geological  point  of  view. 

Twelve  miles  southeast  of  South  Pagi  there  is  a  small  coconut-cov- 
ered island  called  Sanding  which  apparently  is  little  more  than  a  wide 
exposed  reef.  It  has  never  been  visited  by  a  scientist.  South  of  Sanding 
the  two  hundred  meter  curve  ends.  Some  fifty-five  miles  southeast 
of  Sipora  there  is  a  small  pinnacle  called  Mega,  two  miles  long  and 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  317 

surrounded  by  a  fringing  reef.  It  is  low  and  densely  wooded  and  rises 
from  the  one  thousand  meter  curve.  This  island  too  has  never  been 
scientifically  explored. 

Enggano,  the  most  isolated  island  of  this  archipelago,  lies  about  one 
hundred  eighty  miles  south  east  of  Sipora.  It  is  about  seventy  miles 
from  the  nearest  part  of  the  Sumatra  coast.  The  island  is  twenty  miles 
long  and  approximately  ten  broad.  A  range  of  hills  runs  from  north- 
west to  southeast  through  the  center  of  the  island,  expressing  thus  the 
direction  of  upthrust  throughout  the  group.  Enggano  lies  entirely  on 
its  own  shelf,  the  surrounding  seas  reaching  a  depth  of  three  thousand 
meters.  Enggano  is  heavily  wooded  and  the  interior  is  so  marshy  and 
difficult  to  penetrate  that  communication  between  the  villages  is  said 
to  be  by  boat.  It  is  sparsely  inhabited,  the  1919  census  having  shown 
only  four  hundred  seven  persons  on  the  island.  The  main  trade  of  the 
island  is  in  copra,  and  the  principal  settlement  is  on  Aduwe,  one  of  the 
three  little  islands  in  Enggano  Bay.   This  group  is  called  Pulu  Dua. 

FAUNAL  LIST  OF  THE  BIRDS 

In  the  following  list  of  the  birds  of  the  west  Sumatra  islands,  the 
arrangement  of  the  families  has  been  that  of  Wetmore  (A  Systematic 
Classification  for  the  Birds  of  the  World,  Smith.  Misc.  Coll.,  99,  7, 
1940,  11  pp.),  while  the  arrangement  of  the  species  within  the  families 
has  been  that  of  Chasen  (1935).  All  measurements  are  in  millimeters, 
the  wing  pressed  flat  against  the  ruler.  All  weights  of  small  birds  are 
in  grams.  Where  used,  wing-tail  ratio  refers  to  the  percentage  of  the 
length  of  the  tail  as  compared  to  that  of  the  wing.  Wing  tip  index 
refers  to  a  similar  percentage  of  the  length  of  the  shortest  primary  as 
compared  to  that  of  the  longest  primary.  Color  notes  follow  Ridgway. 

Of  the  total  of  seventy-five  families  found  on  Sumatra  and  generally 
in  the  vicinity  of  these  islands,  forty-six  families  are  represented  from 
this  archipelago  with  certainty.  Of  the  missing  families  some  members 
of  the  Phaethontidae,  Sulidae,  Rostratulidae,  Indicatoridae,  and  Arta- 
midae,  may  be  expected  to  turn  up  on  these  islands  in  the  future. 
Notable,  however,  is  the  lack  of  representatives  of  the  Megapodiidae 
and  Phasianidae,  which  do  not  occur  there  and  presumably  never  have 
been  found  on  these  islands. 


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Family  SULIDAE,  Boobies,  Gannets 

The  only  record  for  this  family  is  a  dubious  one  which  I  have  not 
included  in  my  definite  list  of  species  of  the  islands.  There  is  every 
reason  to  suppose,  however,  that  boobies  may  occur  around  these 
waters  from  time  to  time. 

?  Sula  sula  rubripes  Gould 

Chasen  (1935)  records  this  form  from  the  Pagi  Islands.  I  can  find 
no  record  of  a  specimen  from  this  area  except  that  in  the  British 
Museum  Catalogue  (Vol.  XXVI,  p.  434,  1898),  which  lists  a  juvenal 
male  from  Nassau  Island,  south  of  Sumatra,  August  (Dr.  Coppinger), 
Voyage  of  H.M.S.  ''Alert".  In  the  report  on  this  voyage  (British 
Museum,  1884),  Dr.  Coppinger  notes  in  the  summary  of  the  trip  (p.  3) 
that  the  boat  touched  at  Nassau  Island  in  the  Danger  Group  northeast 
of  Samoa,  after  leaving  Tahiti  on  August  eighteenth.  There  is  no 
mention  of  a  stop  near  south  Sumatra  during  the  later  part  of  the  trip, 
although  a  stop  was  made  at  Singapore  in  November  of  the  following 
year.  More  evidence  is  needed  to  prove  the  existence  of  this  form  off 
the  west  Sumatran  islands. 


Family  PHALACROCORACIDAE,  Cormorants 

The  only  record  for  this  family  is  from  Nias.  There  is  no  indication 
as  to  whether  or  not  the  Little  Cormorant  is  a  resident  in  the  islands . 

1.  Phalacrocorax  niger  (Vieillot) 

Listed  by  Biittikofer  (1896)  from  Nias  on  the  basisof  Nieuwenhuisen 
and  von  Rosenberg's  report  (1863).  This  species  has  not  since  been 
recorded  from  any  of  the  islands. 

Family  ARDEIDAE,  Herons 

2.  Ardea  sumatrana  sumatrana  Raffles 

Recorded  from  Simalur,  Babi,  Tuangku,  Nias,  and  the  Pagi  Islands. 

The  record  for  the  Pagi  Islands  is  by  Chasen  and  Kloss  (1935). 
I  can  find  no  specimen  on  which  it  is  based.  A  nest  with  two  well- 
grown  young  was  found  by  Abbott  on  Simalur  in  November.  A  female 
adult  in  the  collection  of  the  National  Museum  was  secured  on  the 
same  island,  October  23,  1902.  Wing,  434. 


RIPLEY:    BIRD   FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  319 

3.  Ardea  purpurea  manilensis  Meyen 

Found  on  Xias,  Pini  (Batu  Islands),  and  Enggano. 

An  immature  specimen  was  taken  on  Enggano  by  Modigliani,  May 
19, 1891.  An  adult  female  collected  by  Dr.  Abbott,  November  5, 1904, 
on  the  same  island  was  labeled  "iris  yellow".   Wing,  351. 

4.  Butorides  striatus  spodiogaster  Sharpe 
Synonym:  Butorides  striatus  sipora  Chasen  and  Kloss 

Occurs  on  Simalur,  Nias,  Siberut,  Sipora  and  North  Pagi. 

Three  specimens  from  Simalur,  Nias,  and  Sipora  measure:  wing 
(worn),  c?  182.5,  9  170.5,  o  180.5;  tail,  d*  63,  9  63.5,  o60;  culmen, 
J1  63,  9  60,  o  60. 

Oberholser  (1912,  p.  1)  named  two  races,  actophilus  from  North 
Pagi  Island  and  icastopterus  from  Simalur,  on  the  basis  of  larger  size 
and  darker  color.  The  types  measure : 

actophilus  9  ad.,  icastopterxis c? ad., 

January  4,  1903.  December  10,  1903. 
wing                                                 196  191 

tail  70  68 

culmen  66.5  65.5 

Another  female  from  North  Pagi,  collected  December  31,  is  also 
large  (wing  190,  tail  67.5,  culmen  62.5).  These  birds  presumably 
represent  a  northern  race  found  in  Siam  and  China  which  migrates 
during  the  winter  to  the  southern  islands.  Chasen  and  Kloss  (1926) 
have  described  sipora  as  being  darker  on  the  neck  than  javanicus, 
although  similar  in  size.  The  three  small  specimens  agree  with  this 
description,  as  well  as  appearing  darker  on  the  back  and  scapulars 
than  a  large  series  of  javanicus.  However,  they  are  unfortunately  in- 
separable from  spodiogaster  Sharpe  (Bull.  Brit.  Orn.  CI.,  3,  1894, 
p.  xvii)  of  the  Andamans  and  Nicobars.  A  female  from  Little  Nicobar 
collected  March  3  has  a  wing  of  180  and  agrees  well  in  color  with  the 
west  Sumatra  islands. 

The  names  carcinophilus  and  carcinophonus  Oberholser  (Journ. 
Wash.  Acad.  Sci.,  14,  1924,  p.  294)  and  abbotti  Oberholser  (U.  S.  Nat. 
Mus.  Bull.  159,  1932,  p.  14)  are  synonyms  of  javanicus. 

5.  Butorides  striatus  actophilus  Oberholser 
Synonym:  Butorides  javanicus  icastopterus  Oberholser 

Migrant  from  the  Asiatic  mainland  recorded  from  Simalur  (icastop- 
terus) and  the  Pagi  islands  in  December  and  January. 


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6.  Ardeola  bacchus  (Bonaparte) 

Recorded  by  von  Rosenberg  from  Nias  (Biittikofer,  1896)  and  a 
sight  record  (Richmond,  1903)  for  Simalur. 

7.  Egretta  garzetta  nigripes  (Temminck) 

There  is  a  record  of  this  species  for  Nias  in  Biittikofer 's  list  based 
on  Nieuwenhuisen  and  von  Rosenberg  (1868). 

8.  Egretta  eulophotes  (Swinhoe) 

A  rare  migrant  from  China  where  it  has  been  seriously  reduced  by 
plume  hunters.    Recorded  from  Sipora  by  Chasen  and  Kloss  (1926) 

9.  Demigretta  sacra  sacra  (Gmelin) 

Occurs  on  Simalur,  Tuangku,  Nias,  Pulo  Tello,  Sipora,  the  Pagi 
islands,  and  Enggano. 

Two  specimens  collected  by  Dr.  Abbott  on  Simalur  and  Pulo  Mirbau 
off  Enggano  measure:  9  ad.,  wing  262,  tarsus  68;  c?  im.,  wing  266.5, 
tarsus  69.5.  The  immature  bird  is  in  the  mottled  juvenal  plumage 
for  the  white  phase  discussed  by  Mayr  and  Amadon  (Amer.  Mus. 
Novit.  No.  1144,  Oct.  13, 1941).  Curiously  enough,  the  wing  measure- 
ment of  the  adult  female  is  smaller  than  any  of  the  series  of  three 
hundred  seventy-eight  specimens  measured  by  them. 

Of  the  thirty-three  known  adults  collected  on  these  islands,  nineteen 
are  described  as  being  in  the  grey  phase,  thirteen  in  the  white  phase, 
and  one  (Salvadori,  1892)  in  the  mottled  phase. 

The  reef  heron  probably  nests  on  the  small  islets  off  the  coasts  of 
the  larger  islands.  Junge  (1936)  records  a  set  of  eggs  taken  July  3 
on  an  islet  in  Sinabang  harbor,  Simalur. 

10.  Mesophoyx  intermedia  intermedia  (Wagler) 

The  single  record  for  this  species  is  for  Nias,  Biittikofer  (1896) 
founded  on  von  Rosenberg's  list. 

11.  Nyticorax  nycticorax  nycticorax  (Linnaeus) 

Found  on  Nias  (Biittikofer  1896)  and  Enggano.  Two  males  and 
a  female  from  the  latter  island  measure:  wing,  c?  270,  287,  9  281. 
One  male  is  in  an  erythristic  state  of  plumage  similar  to  that  recorded 
for  N.  n.  hoactli  by  van  Rossem  (Auk,  53,  1936,  p.  322).   The  cheeks 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  321 

and  nape  are  tinted  with  buff.  The  upper  part  of  the  back  is  dark 
bronzy  brown  shading  posteriorly  to  "Carob  brown".  The  wings  and 
tail  are  greyish  buff.  The  second  male  approaches  this  but  is  paler  and 
more  greenish  on  the  back.  The  female  is  nearly  normal.  The  occur- 
rence of  erythrism  in  this  species  indicates  that  Xycticorax  caledonicus 
may  be  a  form  in  which  this  type  of  mutation  has  been  established  as 
a  dominant.  It  will  be  interesting  to  observe  the  apparently  new 
colony  of  caledonicus  breeding  in  East  Java,  (Hoogerwerf,  Bull.  Raffles 
Mus.,  12,  1936,  p.  120),  to  see  if  the  birds  hybridize  with  nyciticorax. 
The  colors  of  the  soft  parts  are  recorded  as  follows :  iris  straw  yellow; 
bill  black,  base  and  lores  greenish  (cT),  lores  slaty  (cf),  and  base 
pinkish  fleshy  (  9  ) ;  feet  pale  greenish  yellow  (d\  9  ),  straw  yellow  (cf ). 

12.    GORSACHIUS   MELANOLOPHUS   MELANOLOPHUS    (Raffles) 

Two  females  collected  by  Dr.  Abbott  on  Enggano  seem  to  be  the 
only  record  for  this  species  in  the  west  Sumatra  islands.  One  specimen, 
collected  November  22,  1904,  is  in  immature  plumage.  The  soft  parts 
are  as  labelled:  "iris  pale  greenish  yellow;  feet  brownish  green;  bill 
black  above,  pale  brown  beneath." 

13.  Ixobrychus  sinensis  sinensis  (Gmelin) 

An  immature  male  collected  by  Abbott  on  Nias,  March  28,  1903,  is 
the  second  record  for  that  island. 

14.  Ixobrychus  cinnamomeus  (Gmelin) 

A  single  male  is  recorded  from  Enggano  June  22,  1936,  by  June 
(1938). 

15.  Dupetor  flavicollis  flavicollis  (Latham) 
Recorded  from  Xias  by  von  Rosenberg. 

Family  CICOXIIDAE,  Storks 

The  first  member  of  this  family  found  on  these  islands  proves  to 
belong  to  a  widely-distributed  form  throughout  the  Malay  area. 

16.  Dissoura  episcopus  stormi  (Blasius) 

North  Pagi.  An  adult  male  and  an  immature  female  taken  on  North 
Pagi  in  December  by  Menden  are  in  the  collection  of  the  Museum  of 


322  bulletin:  museum  of  comparative  zoology 

Comparative  Zoology.  They  are  the  first  specimens  of  this  family  to 
be  recorded  from  the  west  Sumatra  islands.  The  male  specimen  has 
the  following  notes  on  the  colors  of  the  soft  parts :  "iris  brown,  bill  and 
feet  red"  (but  basal  third  of  bill  is  black  in  the  skin).  The  immature 
has  the  colors  noted  as:  "iris  dark  brown,  feet  red,  bill  red-brown" 
(not  black  basally  in  skin). 

I  am  indebted  to  Mr.  James  L.  Peters  for  the  identification  and  the 
information  on  the  labels. 


Family  ANATIDAE,  Ducks  and  Geese 

The  Javan  Tree  Duck  is  the  only  species  recorded  on  the  west 
Sumatra  islands  so  far.  It  is  a  wandering  species  throughout  the 
Greater  Sunda  islands.  Very  likely  Dendrocyqna  arcuata  occurs  here 
from  time  to  time,  while  the  absence  of  Anas  castanea  gibberifrons 
from  this  area  is  definitely  puzzling,  (see  Ripley,  Auk,  59,  1942,  p.  98). 

17.  Dendrocygna  javanica  javanica  (Horsfield) 

Biittikofer  (1896)  lists  this  as  the  probable  race  encountered  by 
von  Rosenberg.  When  I  was  on  Nias  in  June,  1939,  I  was  told  that 
ducks  were  common  on  Nias  at  certain  seasons. 


Family  ACCIPITRIDAE,  Hawks,  Eagles 

Sixteen  members  of  this  family  have  been  recorded  from  these 
islands  of  which  ten  are  presumably  residents  and  six  migrants.  Five 
of  these  resident  forms  are  presumed  to  be  identical  with  those  found 
commonly  throughout  the  Malayan  area.  Of  the  remaining  five, 
Spizaetus  cirrhatus  vanheurni  is  described  as  being  smaller  than  its 
nearest  relative.  The  other  four  forms  belong  to  the  very  plastic 
Spilornis  cheela  "rassenkreis".  The  two  races  from  the  northern 
islands,  abbotti  and  asturinus  resemble  birds  from  the  Andaman  and 
Nicobar  Islands.  The  race  from  the  Batu  Islands  has  been  united 
with  the  population  from  south  Sumatra.  The  southernmost  race, 
that  from  Sipora  and  the  Pagi  islands  has  characters  which  markedly 
resemble  those  found  in  Java,  Bawean  and  Celebes.  In  all  these  Spi- 
lor?iis  forms  there  are  marked  color  differences. 

18.  Pernis  apivorus  ptilorhynchus  (Temminck) 
Siberut  (Chasen  and  Kloss,  1926,  p.  279.). 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA   ISLANDS  323 

19.    HALIASTUR   INDUS   INTERMEDIUS   Blyth 

Found  on  Simalur,  Babi,  Tuangku,  Nias,  and  the  Pagi  islands. 

Junge  (1936)  notes  that  this  bird  was  found  nesting  on  Simalur  in 
May  in  a  coastal  grove  of  casuarinas  at  a  height  of  20  m.  above  the 
ground.  An  immature  bird  was  collected  at  the  spot. 

20.  Accipiter  badius  poliopsis  (Hume) 

An  adult  female  collected  on  November  20  is  recorded  from  Nias 
by  Buttikofer  (1896). 

21.  Accipiter  soloensis  (Horsfield) 

Taken  by  Abbott  on  Simalur  in  October  and  January.  Chasen 
(1935,  p.  71)  lists  this  species  from  Nias  on  the  basis  of  Biittikofer's 
record  of  Accipiter  poliopsis.  From  the  description  of  the  specimen, 
however,  it  is  impossible  to  be  sure  that  he  did  not  have  a  specimen  of 
poliopsis. 

22.  Accipiter  trivirgatus  trivirgatus  (Temminck) 

Recorded  for  Nias.  A  male  collected  March  26  is  in  extremely  worn 
plumage  showing  no  sign  of  new  feathers.  A  female  collected  on  the 
same  date  is  also  very  worn,  but  there  are  a  few  new  feathers  coming 
in  on  the  sides  of  the  neck  and  nape.  Another  female  collected  March 
23  is  renewing  some  of  the  tail  feathers  and  the  inner  primaries.  A 
few  of  the  nape  feathers  and  wing  coverts  are  new  also.  Soft  parts: 
iris,  cf ,  "bright  yellow";  9  "orange  yellow";  feet,  d*  "yellow",  9 
"pale  yellow" ;  bill,  9  "black  above,  leaden  beneath".  The  stomach 
of  the  female  (March  23)  contained  lizards. 

23.  Accipiter  virgatus  gularis  (Temminck  and  Schlegel) 

Simalur,  Siumat,  Lasia,  and  Nias.  Four  immature  females  from 
Simalur  and  Siumat  and  Lasia  measure:  wing,  182,  185,  185.5,  188. 
Two  of  the  specimens  had  been  eating  small  birds.  This  species  has 
been  collected  from  November  25  to  February  12. 

24.  Spizaetus  nipalensis  alboniger  (Blyth) 

Found  on  Pulu  Asu,  off  Simalur,  Bangkaru  (sight),  Nias,  and  North 
Pagi.  Chasen's  record  (1926)  for  S.  cirrhatus  limnaetus  on  the  Pagi 
islands  presumably  referred  to  this  species.    Three  immature  males 


324  bulletin:  museum  of  comparative  zoology 

from  Pulu  Asu,  Nias,  and  North  Pagi  measure:  wing,  296,  308,  310. 
They  were  taken  December  25,  March  13,  January  6.  Soft  parts: 
iris,  "greyish-yellow,  lemon  yellow";  feet,  "dull  lemon  yellow";  claws, 
"black". 

25.  Spizaetus  cirrhatus  vanheurni  Junge 

This  race  was  described  by  Junge  (1936)  on  the  basis  of  five  speci- 
mens from  Simalur.  The  author  notes  that  the  birds  are  much  smaller 
than  S.  c.  limnaeetus,  but  failed  to  compare  them  with  S.  nipalensis 
alboniger.  In  measurements  and  in  coloration,  these  birds  seem  very 
similar  to  alboniger. 

26.  Ictinaetus  malayensis  malayensis  (Temminck) 

Listed  by  Biittikofer  from  Nias  on  the  basis  of  Nieuwenhuisen  and 
von  Rosenberg  (1863).  This  mountain  forest  species  has  never  been 
encountered  since  on  the  west  Sumatra  islands. 

27.  Haliaeetus  leucogaster  (Gmelin) 

A  bird  of  the  small  islands  off  Simalur;  Asu,  Djawi  djawi,  Babi- 
ketchil,  all  in  Sinabang  bay,  Siumat,  and  Lasia,  Babi,  Bangkaru, 
Tuangku,  Nias  (vide  Chasen,  1935),  Sipora,  and  the  Pagi  islands. 

Two  males  were  collected  on  Asu  and  Siumat  in  December.  A  nest 
was  seen  by  Abbott  on  Siumat  on  Christmas  Daj^. 

28.  Spilornis  cheela  abbotti  Richmond 

Simalur.  This  well-marked  race  was  collected  by  Abbott  (1901)  and 
van  Heurn  (1913)  at  all  months  of  the  year.  The  wings  of  nine  males, 
six  females,  and  one  unsexed,  measure:  cfcf  330-362,  9  9  319-359, 
o328. 

Soft  parts:  "iris  yellow;  bill  horn  blue,  brownish  at  tip;  cere  orange 
yellow;  inside  of  mouth  leaden;  feet  dirty  yellow." 

Stomach  contents:  snakes,  small  crab,  centipede,  lizards. 

Apparently  an  abundant  bird  on  Simalur,  this  race  is  very  distinc- 
tive in  appearance.  The  amount  of  spotting  on  the  under  parts  is 
variable.  In  some  specimens  it  extends  up  onto  the  breast;  in  others  it 
is  confined  to  the  abdomen.  The  marking  on  the  breast  and  throat  is 
far  more  noticeable  than  in  the  paler  forms,  due  to  the  contrast  be- 
tween the  dark  blackish  brown  and  reddish  brown  bars. 

No  pale  individuals  have  so  far  been  taken.  Meise's  recent  revision 
of  this  genus  (J.f.O.,  87,  1, 1939,  p.  65)  indicates  that  all  these  popula- 
tions should  be  considered  races  of  cheela. 


RIPLEY:    BIRD   FAUNA    OF   THE    WEST   SUMATRA    ISLANDS  325 

29.  Spilornis  cheela  asturinus  A.  B.  Meyer 

Nias.  Three  males  collected  by  Abbott  measure:  wing,  291,  291.5, 
299.5  All  three  birds  are  in  worn  plumage  with  a  few  new  feathers 
appearing  among  the  greater  wing  coverts  (one  male)  and  secondaries 
(another  specimen).  Soft  parts:  iris  bright  yellow,  feet  dirty  orange 
yellow,  cere  yellow.   Stomach  contents:  snakes  and  small  centipedes. 

A  pair  collected  by  myself  for  the  Academy  of  Natural  Sciences 
weighed,  d*  15  oz.,  9  1  lb.  4  oz. 

This  species  is  common  on  Nias,  where  it  may  be  found  perching  in 
a  high  tree  on  the  edge  of  an  open  field  or  moving  slowly  over  the 
gardens  or  hedgerows  in  search  of  its  prey. 

From  minimus  of  the  Northern  Nicobars,  which  it  closely  re- 
sembles, this  form  differs  by  the  finer  barring  and  spotting  of  the 
lower  underparts. 

30.  Spilornis  cheela  batu  deSchauensee  and  Ripley 

Tello,  Tana  Massa.  Meise  (I.e.  1939)  in  his  splendid  revision  of 
the  genus  Spilornis  lumps  Malay  Peninsula,  Sumatra,  Java,  and  Bali 
birds  all  under  bassus,  giving  for  wing  measurements,  348-394.  I  am 
inclined  to  believe  that  the  name  bido  should  stand  for  Java  and  Bali 
birds,  which  are  large  (384,  391),  and  dark,  with  distinctly  blackish 
throats.  Granted  the  last  character  is  allelic  in  these  birds,  still  it 
seems  to  be  quite  constant  in  the  Java  population. 

The  birds  from  south  Sumatra,  on  the  other  hand,  are  uniformly 
small,  distinctly  paler  in  tone  than  bido,  although  in  this  last  character, 
they  do  not  differ  from  typical  bassus. 

When  de  Schauensee  and  I  described  batu  (1940,  p.  401.),  we  had 
only  north  Sumatran  material  available  for  comparison.  An  adult 
from  Tana  Massa  in  the  National  Museum's  collection  (sex  indet. 
no.  179622)  does  not  bear  out  our  contention  that  this  is  a  pale  race. 
It  is  as  dark  in  tone  as  any  specimen  from  Sumatra  or  the  Malay 
Peninsula.  However,  it  does  agree  with  the  Academy's  two  birds  in 
being  small.  Four  males,  three  females,  and  one  sex  indet.  from  the 
east  coast  province  of  Sumatra  agree  with  these  Batu  Island  birds  in 
size  and  color.  They  come  from  the  Katenan  River,  the  Siak  and 
Little  Siak  rivers,  Makapan,  and  Pulu  Padang.  Their  measurements 
are:  <?d\  wing  229,  331,  346.5,  352,  tail  204,  206,  217  (molt),  220; 
9  9  ,  wing  340,  350,  354,  tail  218,  221,  221 ;  o  wing  346,  tail  215. 

As  can  be  seen,  these  measurements  are  all  below  360  for  the  wing 
and  230  for  the  tail.  North  Sumatra  birds,  on  the  other  hand,  measure 


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much  larger.  Six  males  and  four  females  from  Atjeh  measure:  wing, 
c?  360,  365*,  365,  388.5,  390*,  410.5;  9  371*,  375,  380,  388;  tail,  <?  231 
255*;  9  253*.  Similarly  seven  Malay  Peninsula  birds  measure  wing, 
cf  355  (1,  worn)  -  397  (374);  9  385^07  (392);  &  230-259  (244.6), 
9  242-264  (251.6). 

Under  these  circumstances,  I  prefer  to  restrict  the  name  bassus  to 
birds  from  the  Malay  Peninsula  and  Sumatra  south  to  the  province 
of  Oostkust.  I  propose  also  to  unite  birds  from  Oostkust  south  with 
Batu  Island  birds  as  batu,  modifying  the  description  of  that  race 
(I.e.  1940,)  to  read:  Pulo  Tello  and  South  Sumatra  birds  "differ  by  .  .  . 
smaller  size". 

31.  Spilornis  cheela  sipora  Chasen  and  Kloss 

Sipora  and  the  Pagi  islands.  A  single  female  from  Sipora  has  a 
wing  measurement  of  310  and  the  soft  parts  are  indicated  as:  "iris 
and  facial  skin  yellow;  bill  pale  grey;  feet  yellow". 

This  race  is  distinctive  in  having  the  feathers  of  the  crest  largely 
white  with  black  tips  giving  a  noticeably  piebald  effect.  The  breast 
is  solidly  colored,  smoky  bronzy  black.  In  its  coloration,  sipora  shows 
a  close  relationship  to  the  black-cheeked  strain  of  Spilornis  which  con- 
tinues southward  with  bido  of  Java,  baweanus  of  Bawean,  and  rufipectus 
of  Celebes. 

Family  FALCONIDAE  Falcons 
Two  species  have  been  recorded  from  the  islands,  both  migrants. 

32.  Falco  peregrinus  calidus  Latham 

A  migrant  collected  on  Nias  and  Enggano  and  probably  seen  on 
Simalur  (Richmond,  1903).  A  male  from  Nias  collected  in  March  has 
a  wing  of  306.5.  A  female  from  Enggano  collected  in  November 
measures  376.  The  latter  specimen  is  molting  the  greater  wing  coverts. 
The  soft  parts  of  the  male  are  given  as:  "iris  dark  brown;  eyelids 
yellowish  grey;  feet  bright  yellow". 

33.  Falco  tinnunculus  subsp.  ? 

A  winter  migrant  recorded  from  Nias  in  November  by  Biittikofer 

(1896). 

•  U.S.N.M.  birds. 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  327 


Family  RALLIDAE,  Rails,  Coots,  Gallinules 

Four  species  occur  on  the  west  Sumatra  islands  all  apparently 
identical  with  wide-ranging  Malayan  forms. 

34.  Rallus  striatus  gularis  Horsfield 
Synonym:  Hypotaenidia  striata  reliqua  Oberholser 

Simalur.  Two  females  collected  in  October  and  December  are 
identical  with  specimens  from  Java  and  Singapore.  The  soft  parts  are 
marked  as:  "iris  pale  yellow  brown,  red;  bill  pinkish  red,  tip  horn 
brown;  feet  dull  brownish  purple".  Found  in  the  rice  paddies. 

(Rallus  jouyi  Stejneger  is  by  no  means  a  synonym  of  gularis.  It  is 
a  giant  pale  race.  The  male  type  and  a  female  from  Shanghai  measure : 
wing,  cf  135,  9  134.5;  tail,  cf  50,  9  49;  culmen,  d"  43.5,  9  42.5.) 

35.  Rallina  fasciata  (Raffles) 

Nias,  Sipora,  and  Enggano.  A  male  from  Enggano  collected  in 
November  measures:  wing  128.5.  Soft  parts:  "iris  and  eyelids  red;  feet 
vermilion." 

36.  Amaurornis  phoenicurus  javanica  (Horsfield) 
Synonym:  Amaurornis  phoenicura  cleptea  Oberholser 

Found  on  Simalur,  Nias,  Tello,  Siberut,  Sipora,  the  Pagi  islands,  and 
Enggano. 

A  common  bird  of  the  flooded  rice  fields  and  swampy  meadows  near 
streams.  Found  also  in  the  sago  swamps  on  those  islands  which  have 
them.  These  birds  have  a  harsh  rattling  call. 

Four  males  from  Simalur,  three  females  from  Nias,  including  the 
type  of  cleptea,  and  four  females  from  Sipora  are  inseparable  on  the 
basis  of  size. 

37.  Porphyrio  poliocephalus  indicus  Horsfield 
Recorded  from  Nias  by  von  Rosenberg  (1878). 


Family  CHARADRIIDAE,  Plovers 

Four  species  have  been  found  on  these  islands  of  which  three  are 
migrants.  The  resident  form  has  been  separated  on  the  basis  of  color, 
a  character  which  I  fail  to  distinguish. 


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38.  Pluvialis  dominica  fulva  (Gmelin) 

A  common  migrant  recorded  from  Simalur,  Nias,  Batu  islands, 
Siberut,  and  Sipora,  September  through  April. 

39.  Charadrius  peronii  Schlegel 
Synonym:  Charadrius  peronii  chaseni  Junge 

A  resident  species  recorded  from  Simalur  and  Nias.  A  female  from 
Simalur  (M.C.Z.  178151)  was  collected  in  May.  Junge  described  the 
race  chaseni  from  Simalur  as  being  "darker  on  the  upper  parts,  less 
rufous  on  the  head  especially  in  the  9  9  .  Also  the  rufous  colour 
behind  the  white  neckcollar  and  on  the  sides  of  the  breast  is  much  more 
pronounced  than  in  p.  peronii.  The  bill  in  chaseni  is  slightly  heavier." 

A  female  from  Nias  collected  in  February  (two  months  earlier  than 
Junge's  Simalur  series)  is  a  good  deal  darker  than  the  Simalur  female. 
It  is  in  much  fresher  plumage.  A  molting  female  from  the  Philippines 
collected  in  January  has  new  feathers  as  dark  as  the  Nias  bird,  while 
an  October  male  from  Mindanao  is  considerably  darker  than  any  other 
bird  in  our  series.  As  far  as  the  character  of  the  rufous  breast  band 
goes,  the  two  west  Sumatra  island  birds  stand  midway  in  a  series  of 
seven  females.  I  cannot  find  any  significant  size  difference  in  the  bill. 
I  fail  to  see,  therefore,  how  chaseni  can  be  upheld. 

40.  Charadrius  mongolus  atrifrons  Wagler 

A  migrant  recorded  from  Simalur  and  Tello;  this  species  probably 
occurs  on  all  the  small  beaches.  Two  females  from  Simalur  collected 
in  December  measure:  wing,  130,  134.5. 

41.  Charadrius  leschenaultii  Lesson 

A  winter  migrant.  There  are  records  for  Simalur,  Nias,  Pini,  Tello, 
and  Sipora  from  August  through  December.  A  male  and  two  females 
collected  on  Simalur  in  December  measure:  wing,  cf  140,  9  140.5, 
142.5. 

Family  SCOLOPACIDAE,  Snipe,  Woodcock,  Sandpipers 
Twelve  species  have  been  found  on  these  islands,  all  of  them  migrants. 

42.  Numenius  phaeopus  phaeopus  (Linnaeus) 

A  migrant  taken  on  Simalur,  Sipora,  and  Siberut.  A  female  taken 
on  Simalur  in  November  has  the  rump  pure  white.    A  male  from 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  329 

Sipora  collected  in  February  has  a  few  spots  of  grey  brown  among  the 
white  rump  feathers.  For  the  time  being,  I  think  it  is  better  to  call 
these  birds  phacopus.  There  seems  to  be  a  good  deal  of  intergradation 
between  the  two  forms. 

43.    NUMENIUS  PHAEOPUS  VARIEGATUS  (Scopoli) 

Reported  from  August  till  June  on  Simalur,  Nias,  Pini,  and  Tello. 
There  is  a  sight  record  of  one  or  other  of  the  races  of  whimbrel  for  the 
Pagi  islands. 

44.  Numenius  arquatus  orientalis  C.  L.  Brehm 

A  migrant  taken  on  Tuangku  and  Nias.  A  female  from  Tuangku 
(January)  measures:  wing,  282. 

45.  Limosa  lapponica  baueri  Xaumann 

The  single  record  of  this  migrant  for  the  west  Sumatra  islands  is 
Xias,  October  (Blasius). 

46.  Tringa  totanus  eurhinus  (Oberholser) 

This  winter  visitor  has  been  taken  on  Simalur,  Nias,  and  Tana 
Massa  from  September  until  May.  A  female  from  Nias  collected  in 
March  has  a  wing  measurement  of  158. 

47.  Tringa  nebularia  Gunnerus 

A  migrant  reported  from  Simalur  and  Nias  in  August,  September, 
and  October. 

48.  Tringa  glareola  Linnaeus 

This  winter  bird  was  found  on  Simalur  from  January  until  April. 

49.  Actitis  hypoleucos  (Linnaeus) 

The  common  sandpiper  of  the  islands  from  August  until  April. 
Recorded  with  certainty  only  from  Simalur,  Babi,  Nias,  Tello,  and  the 
Pagi  islands. 

50.  Arenaria  interpres  interpres  (Linnaeus) 

A  female  collected  on  Simalur  in  December  has  a  wing  measurement 
of  148.5. 


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51.  Capella  stenura  (Bonaparte) 

During  migration  recorded  from  Simalur,  Nias,  Sipora,  and  (sight 
record)  the  Pagi  islands.  Two  males  and  three  females  from  Simalur 
and  Nias  measure  :wing,  &  135,  138;  9  134.5, 135, 135.5.  These  birds 
were  collected  during  December,  February,  and  March. 

The  Pintail  Snipe  is  a  common  bird  in  these  islands  during  the 
winter,  being  found  wherever  there  are  recently  drained  rice  fields  or 
along  the  edges  of  streams,  sometimes  in  the  short  grass  bordering 
roads  built  through  swampy  areas. 

52.  Erolia  ruficollis  (Pallas) 

A  casual  winter  visitor,  found  on  Simalur  and  Nias  in  December 
and  October. 

53.  Erolia  testacea  (Pallas) 
Blasius  (1901)  records  this  migrant  as  taken  on  Nias  in  October. 


Family  BURHINIDAE,  Thick-knees 
A  single  resident  species  occurs  throughout  the  greater  Sunda  area. 

54.  Orthorhamphus  magnirostris  (Vieillot) 

Simalur,  Babi,  Tuangku,  and  Nias.  A  male  from  Babi  collected  in 
January  has  a  wing  measurement  of  291.5.  Weight,  2.5  pounds.  Soft 
parts:  "iris  yellow,  tarsi  pale  yellowish  slaty,  thighs  yellow,  claws 
black,  leaden  at  base".  The  Stone-plover  feeds  on  crabs  and  is  common 
along  sandy  beaches  and  on  exposed  reefs  at  all  seasons. 


Family  GLAREOLIDAE,  Pratincoles 
A  single  species  of  this  family  has  been  recorded  twice  as  a  migrant. 

55.  Glareola  maldivarum  Forster 
A  migrant  from  northern  Asia  recorded  from  Nias  and  Sipora. 


RIPLEY:   BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  331 


Family  LARIDAE,  Gulls,  Terns 

Five  species  have  been  noted  as  occurring  on  these  islands.  At  least 
one  species  is  known  to  breed  here,  but  there  has  been  no  observable 
speciation  in  the  case  of  this  family. 

56.  Chlidonias  hybrida  javanica  (Horsfield) 

There  is  a  record  for  Nias  in  December  (Blasius,  1901),  and  von 
Rosenberg  (1878)  includes  this  in  his  list  of  species  met  with  from  time 
to  time  among  the  islands. 

57.  Sterna  dougallii  bangsi  Mathews 
Collected  on  Siumat  in  May. 

58.  Sterna  sumatrana  sumatrana  Raffles 

Simalur,  Pandjang,  Djawi-djawi,  Siumat,  Nias,  and  Enggano.  Eggs 
have  been  taken  in  May  on  Simalur  and  Djawi-djawi.  An  immature 
male  was  taken  off  Nias  by  Abbott,  October  30. 

59.  Thalasseus  bergii  cristatus  (Stephens) 

Mentioned  by  von  Rosenberg  (1878)  as  occurring  in  the  seas  about 
these  islands. 

60.  Anous  stolidus  pileatus  (Scopoli) 

Listed  by  von  Rosenberg  as  seen  from  time  to  time  among  the 
islands. 


Family  COLUMBIDAE,  Pigeons,  Doves 

Twenty-one  forms  are  recorded  from  the  west  Sumatra  islands.  Of 
these  seven  belong  to  wide-spread  species,  common  throughout  the 
Greater  Sunda  area.  Thirteen  of  the  fourteen  other  forms  are  endemic. 
One  is  considered  identical  with  Sumatran  and  Malayan  subspecies. 
Of  the  thirteen  endemic  races  eleven  differ  markedly  from  their  cor- 
responding relative  on  Sumatra  by  larger  size  and  rather  different 
coloring.  Two  forms  do  not  differ  in  size.  One,  Treron  fulvicollis 
melopogenys,  is  poorly  characterized  and  may  not  be  a  good  race.  The 
other,  Ducula  aenea  consobrina,  differs  from  D.  aenea  only  in  color. 


332 


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The  eleven  well  characterized  forms  may  be  tabulated  as  follows: 


D.  aenea         M .  ruficeps  M.  phasianella 


T.  curvirostra 


En 


ggano* 


zzm 


Mentawi 
&  Paei. 


Xias 


Simalur 


BBZ2Z 
i     I i     i 


Sumatra 


1.    2.    3.    4.     5.     1.    2.    3.     1.    2.    3.  4.    5.    1.    -2.    3.    1.    2.    3.   4.    5. 


M       H    difference  in  size. 
V/J///A   difference  in  color. 


Fig.  3 


In  the  above  chart  Sumatra  birds  are  taken  as  standard.  Their  in- 
dividual variation  is  taken  as  less  than  the  arbitrary  unit  1,  within 
which  it  is  considered  that  local  island  populations  may  vary  and  still 
be  considered  inseparable.  The  unit  2  is  used  to  express  those  varia- 
tions in  size  or  color,  either  one  of  which  by  itself  is  still  too  slight  for 
separation. 

Above  this  number  the  units  used  indicate  the  relative  degree  of 
differentiation  of  the  two  factors,  size  and  color,  of  the  different  island 
populations. 


61.  Treron  curvirostra  haliploa  Oberholser 

Simalur.  The  type,  an  adult  male,  measures:  wing  145.5;  tail  92.5; 
culmen  16.5.  The  soft  parts  are  marked:  iris  orange,  orbital  skin  yel- 
lowish green,  feet  dull  purple.  Junge  (1936)  gives  the  measurements 
of  four  males  as :  wing  146-151 ;  tail  90-94;  culmen  16-17.5.  This  race 
differs  from  curvirostra  by  larger  size.  Thirteen  males  from  Peninsular 
Siam  and  Sumatra  (harterti  is  a  synonym)  measure:  wing  132-139; 
tail  77-83.5;  culmen  16-18. 

Junge  (I.e.)  notes  that  a  single  male  from  Babi  is  quite  yellowish  on 
the  under  surface.  His  description  sounds  very  much  like  smicra  from 
farther  south  (see  also  Junge,  1938,  p.  340.). 

A  bird  of  original  forest. 


RIPLEY:    BIRD   FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  333 

62.  Treron  curvirostra  pega  Oberholser 

Nias.  The  type,  an  adult  male,  measures :  wing  144;  tail  88;  eulmen 
16.  Four  females  measure:  wing  134.5,  142,  143.5,  143.5;  tail  75,  79, 
79,  82;  eulmen  16.5,  17,  17.5,  18.  This  form  is  larger  than  curvirostra 
and  differs  both  from  that  form  and  haliploa  by  being  noticeably  paler 
on  the  under  parts  and  by  having  more  grey  on  the  flanks.  The  soft 
parts  are  recorded  as:  "iris  bright  ochreous ;  orbital  skin  apple  green, 
pale  bluish  green;  feet  maroon  red;  bill  yellow;  cere  red."  Weight: 
cf,  7  oz. 

The  female  differs  as  the  male. 

Specimens  with  enlarged  gonads  were  taken  by  myself  in  June. 

63.  Treron  curvirostra  smicra  Oberholser 

Tello,  Tana  Bala,  Siberut,  and  Sipora. 

The  type,  an  immature  male  taken  February  8  on  Tana  Bala,  meas- 
ures: wing  138.5;  tail  73;  eulmen  17.  Four  males  and  a  female  from 
Siberut  and  Sipora  measure:  wing  d"  141,  141.5,  145,  147.5,  9  148.5; 
tail  d\  83.5,  84,  90,  9  86;  eulmen  &  17.5,  17.5,  17.5,  18,  9  18.  Soft 
parts:  "iris  yellow;  bill  pale  green;  cere  olive;  feet  maroon  red,  crimson 
lake". 

This  race  is  very  bright  yellow  on  the  under  surface.  The  breast 
particularly  is  a  rather  rich  shade  of  lemon  yellow.  It  is  the  brightest 
member  of  the  species.  On  the  upper  surface  it  differs  from  its  near 
relatives  by  having  the  nape  more  distinctly  yellowish  green  less 
greyish  green. 

The  female  differs  as  the  male. 

Evidently  this  bird  prefers  a  slightly  different  type  of  habitat  than 
either  haliploa  or  pega,  which  are  found  on  much  larger  islands.  If 
the  specimen  from  Babi,  one  hundred  and  sixty  miles  to  the  north, 
recorded  by  Junge,  proves  to  belong  to  this  race,  it  would  seem  to 
indicate  that  smicra  not  only  prefers  a  different  ecological  setting  but 
also  suits  its  behavior  to  its  location.  Small  island  populations  of 
pigeons  seem  to  be  more  subject  to  sporadic  wandering  movements 
than  their  large  island  congeners. 

64.  Treron  curvirostra  hypothapsina  Oberholser 

Enggano.  The  male  type,  collected  November  21,  measures:  wing 
156;  tail  92;  eulmen  17.  The  soft  parts  are  recorded  as  follows:  "iris 
vellow;  eyelids  dull  green;  bill  jade  green,  base  dull  green;  feet  purplish 
red." 


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Two  other  males  and  a  female  measure:  wing  c?  149,  150,  9  141; 
tail  cf  86,  90.5,  9  85.5;  culmen  c?  18,  18.5,  9  16. 

This  again  is  a  larger  bird  than  curvirostra  and  much  brighter  yellow 
on  the  under  surface.  It  differs  from  smicra  by  being  slightly  duller  on 
the  under  parts,  lacking  especially  the  bright  suffusion  of  yellow  on  the 
breast  found  in  the  latter  subspecies.  On  the  upper  surface,  too,  the 
nape  is  more  grayish-green,  less  tinged  with  yellow.  Junge  (1938) 
records  enlarged  gonads  in  males  collected  in  June  and  July. 

65.  Treron  fulvicollis  melopogenys  (Oberholser) 

Nias.  The  type  and  only  available  specimen  is  a  female  collected  by 
Dr.  Abbott,  March  18,  1903.  It  measures:  wing  141 ;  tail  85.5;  culmen 

16.  Soft  parts:  "iris  pinkish  mauve;  bill  greenish  blue;  cere  dull 
crimson,  eyelid  gray  with  yellow  edge;  feet  maroon  red." 

Two  females  of  fulvicollis  from  Sumatra  and  Sarawak  measure:  wing 
145,  150.  This  is  a  rather  variable  species  particularly  in  color.  Lack- 
ing a  larger  series,  however,  I  suppose  that  this  single  rather  small 
bright-yellow-throated  specimen  must  be  recognized. 

66.  Treron  olax  (Temminck) 

Recorded  by  von  Rosenberg  (1878)  as  being  on  "all  the  larger 
islands." 

67.  Treron  vernans  miza  (Oberholser) 

Simalur.   The  type,  an  adult  female,  collected  November  22,  meas- 
ures: wing  152.5;  tail  95;  culmen  15.  Soft  parts:  "feet  deep  red." 
Three  males  measure:  wing  156, 156.5, 158;  tail  98,  98.5, 104;  culmen 

17,  17. 

This  is  purely  a  size  race.  There  are  no  color  differences  compared 
to  griseicapilla.  Junge  (1936)  describes  the  plumage  of  a  juvenal  bird 
collected  in  July.    Eggs  were  collected  in  April,  June,  and  July. 

68.  Treron  vernans  griseicapilla  Schlegel 
Synonym:  Dendrophassa  vernans  mesochloa  Oberholser 
Synonym:  Dendrophassa  vernans  polioptila  Oberholser 

Nias,  Tana  Massa,  Tana  Bala,  Siberut,  Sipora,  North  and  South 
Pagi,  Enggano.  The  type  of  mesochloa,  an  adult  female  taken  on  Nias 
March  18,  measures:  wing  145;  tail  81.5;  culmen  16.5.  Soft  parts  are 
recorded  as :  "iris  pinkish-yellow;  bill  gray;  cere  dull  green;  feet  maroon 


RIPLEY:   BIRD   FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  335 

red."  A  series  of  males  and  females  from  Nias  measure:  wing  c?  141 
155,  9  141,  146. 

The  type  of  polioptila,  an  adult  female  collected  on  North  Pagi 
January  1,  measures:  wing  148;  tail  S9.5;  culmen  14.5.  A  series  of 
males  and  females  from  the  Batu  Islands,  Sipora,  North  and  South 
Pagi  and  Enggano,  measure:  wing  cf  148-156,  9   150-152. 

These  birds  differ  a  good  deal  in  size.  Nias  birds  are  more  nearly  the 
size  of  Sumatran  and  Malay  Peninsula  birds,  while  specimens  from 
the  islands  slightly  to  the  south,  particularly  North  Pagi,  tend  to  be 
larger.  In  view  of  the  overlapping  measurements,  however,  it  seems 
to  be  much  wiser  to  group  these  birds  all  under  griseicapilla.  None 
of  the  local  populations  seem  to  be  entitled  to  subspecific  recognition, 
either  on  the  basis  of  standard  deviation  tabulation  or  of  zoogeography. 
Only  the  Simalur  birds  are  apparently  significantly  larger.  It  is  in- 
teresting to  note  that  a  large  series  of  griseicapilla  from  Sumatra  and 
the  Malay  Peninsula  preserve  far  more  uniformity  in  their  wing 
measurements  (cf  144-151,  9  142-150)  than  do  the  birds  from  the 
western  islands.  Presumably  this  is  due  to  less  possibility  of  swamping 
of  the  island  populations. 

Junge  (1938)  records  enlarged  gonads  in  a  male  from  Enggano  taken 
in  June.  This  is  a  common  bird  in  the  islands.  It  is  fond  of  thick 
secondary  growth  in  swampy  coastal  locations.  As  with  all  pigeons, 
they  are  very  fond  of  the  fruit  of  various  species  of  Ficus. 

69.  Leucotreron  jambu  (Gmelin) 

Recorded  from  Nias  in  August  (Blasius)  and  from  the  same  island 
by  Hartert. 

70.  Ducula  aenea  consobrina  (Salvadori) 

Synonym:  Carpophaga  Vandepolli  Biittikofer 

Synonym:  Muscadivores  consobrina  babiensis  Richmond 

Synonym:  Muscadivores  aeneus  mistus  Oberholser 

Synonym:  Muscadivores  aeneus  vicinus  Riley 

Simalur,  Lasia,  Babi,  Tuangku,  Nias,  Tello,  Tana  Bala,  Siberut, 
Sipora,  North  and  South  Pagi. 

The  name  vandepolli  was  founded  on  a  discolored  Nias  bird  (Junge, 
1935). 

The  type  of  babiensis,  an  adult  male  from  Babi,  measures:  wing 
248;  tail  146.5;  culmen,  23.  Two  pairs  from  Lasia  and  Babi  measure: 


336  bulletin:  museum  of  comparative  zoology 

wing,  cf,  245,  247  (246.6);  9  237,  237;  tail,  <?,  144,  146.5;  culmen, 
<?,  24,  26.5. 

The  race  mistus  from  Simalur  was  described  as  being  decidedly 
smaller  than  Nias  birds.  The  type  measures:  wing,  231;  tail,  135.5; 
culmen,  22.5.  The  wing  measurements  of  a  series  of  males  from 
Simalur,  225.5-238  (231.1),  as  against  a  series  from  Nias,  230.5-244 
(235.8),  show  far  too  much  overlap  to  have  any  significance. 

When  the  wing  measurements  of  the  three  males  of  babiensis  are 
compared  with  those  of  twenty-five  males  from  Simalur,  Tuangku, 
Nias,  Tana  Bala,  Sipora,  North  and  South  Pagi,  there  is  only  a  small 
amount  of  overlap  (2.5) : 

wing  average 

adult  c?  d\  Babi  and  Lasia  245-248  (246.6) 

"     other  islands  226.5-247.5  (234.6) 

However,  the  range  of  variation  is  very  great  and  accordingly  I  have 
tested  them  by  the  formula  of  t  used  in  the  comparison  of  the  means 
of  two  small  samples,  (Simpson  and  Roe  "Quantitative  Zoology", 
1939,  p.  210).  Using  this  I  obtained  a  value  for  /  on  the  wing  measure- 
ments of  1.9.  This  gives  a  probability  of  between  .1  and  .05  showing 
that  the  deviation  is  not  significant.  This  bears  out  the  evidence  of 
a  male  from  Babi  recorded  by  Junge  (1936)  which  measured  well 
within  the  range  of  consobrina. 

The  race  vicinus  from  the  Batu  and  Mentawi  islands  was  founded 
on  color:  "the  breast  and  hind  neck  washed  with  much  deeper 
vinaceous-lilac."  The  color  of  these  birds  is  subject  to  much  variation 
through  dirt  or  grease  on  the  feathers.  When  absolutely  clean  speci- 
mens of  each  island  are  compared,  however,  color  variations  tend  to 
disappear. 

From  aenea  these  birds  differ  by  lacking  the  pronounced  vinaceous 
wash  on  the  head,  cheeks,  and  sides  of  the  neck.  As  a  result,  there  is 
less  contrast  between  the  gray  head  and  the  white  margin  around  the 
bill.  The  under  parts  generally  tend  to  be  more  gray,  less  tinged  with 
vinaceous,  although  the  latter  tint  is  noticeable  in  dirty  specimens. 
The  wider  tail  coverts  seem  to  average  darker  in  consobrina.  The 
measuring  of  wings  in  these  birds  is  sometimes  quite  difficult,  due  to 
the  wing  being  incompletely  pulled  back  by  the  skinners.  This  fact 
undoubtedly  accounts  for  some  of  the  variations  in  measurements. 

Molting  specimens  were  collected  in  November  and  immature  birds 
in  October,  November,  and  January.  Eggs  have  been  taken  on  Simalur 
in  May  and  June.    Immature  birds  are  distinguished  by  a  less  well  de- 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  337 

fined  division  between  the  gray  of  the  nape  and  the  bronzy  green  of 
the  back.  Also  the  marginal  under  wing  coverts  are  edged  with 
rufous. 

Soft  parts  of  adults  are  given  as:  "iris,  deep  red;  bill,  pale  leaden, 
base  dark  leaden;  cere,  dull  purple;  feet,  livia  purple."  Immature 
birds  are  given  as:  "iris,  sepia;  bill,  gray;  cere,  faintly  brown;  eyelids, 
gray  edged  with  pale  pinkish  brown;  feet,  purplish  brown." 

These  big  pigeons  are  common  and  conspicuous  on  most  of  the 
islands,  although  on  Nias  they  seem  to  be  confined  to  the  higher  parts 
of  the  island  where  some  original  forest  is  still  found.  The  birds  are 
trapped  a  great  deal  by  the  people  either  as  young  taken  from  the 
nest  or  by  the  use  of  bird  lime  (Malay  "gutta").  Pigeons  are  a  staple 
of  the  native  diet. 

71.  Ducula  aenea  oexothorax  (Salvadori) 

Enggano.  Ten  males  and  two  females  measure:  wing,  d",  247-254; 
9  ,  246,  249;  tail,  d\  152-156;  culmen,  d\  21.5-25.5. 

This  race  differs  from  aenea  by  larger  size  and  by  lacking  the  vina- 
ceous  wash  on  the  head.  In  most  specimens  the  tail  feathers  are  dis- 
tinctly greenish  instead  of  bluish  iridescent  in  color.  On  the  under 
surface,  however,  there  is  an  ill-defined  band  of  strong  vinaceous  wash 
on  the  breast.   The  under  tail  coverts  are  dull  brownish  green. 

From  consobrina  this  race  differs  by  larger  size  and  a  slightly  longer 
tail  proportionate  to  the  length  of  the  wing  (62%  compared  to  57- 
59%).  The  same  characters  of  vinaceous  breast  band  and  more  green- 
ish under  tail  coverts  apply  in  this  case  as  well  as  with  aenea. 

Soft  parts:  iris,  deep  crimson;  bill,  pale  leaden,  base  dark  leaden; 
cere,  dull  purple;  feet,  dull  purplish  red. 

72.  Ducula  bicolor  bicolor  (Scopoli) 

Simalur,  Lasia,  Babi,  Tuangku,  Nias,  Tello,  Lago,  Pini,  Sipora,  the 
Pagi  islands,  and  Enggano.  Specimens  from  Simalur,  Babi,  Sipora, 
and  Enggano  collected  in  October,  November,  December,  and  Jan- 
uary measure:  wing,  d\  224-238,  9  ,  223.5-231.5.  An  immature  bird 
was  taken  on  Sipora  in  October.  The  soft  parts  are  recorded  as:  iris 
dark;  bill  leaden,  black  at  tip;  cere  greenish;  feet  blue. 

This  is  a  bird  primarily  of  the  very  small  islands  and  reefs.  They 
follow  the  seasons  flying  about  in  small  flocks  in  search  of  trees  in 
fruit. 


338  bulletin:  museum  of  comparative  zoology 

73.  Ducula  bicolor  badia  (Raffles) 

Rosenberg  (1878)  records  encountering  this  bird  on  Nias,  the 
Mentawi  islands,  and  Enggano.  It  has  not  been  met  with  since  that 
time. 

74.  Columba  Argentina  Bonaparte 

Recorded  from  Simalur,  Sipora,  and  South  Pagi.  There  is  a  sight 
record  for  Kokos.  Males  and  females  from  Simalur  and  South  Pagi 
measure :  wing,  d\  239-249.5;  9  227,  240.  The  soft  parts  are  given  as: 
iris,  bright  red,  yellow  suffused  with  red;  bill,  greenish  horny,  dull 
purple  at  base,  apple  green;  cere,  dull  purple;  tarsi,  purple;  toes,  pale 
purplish  fleshy. 

This  is  a  wandering  seasonal  species  not  common  in  collections.  It 
probably  occurs  from  time  to  time  on  all  the  west  Sumatra  islands. 

75.  Macropygia  ruficeps  simalurensis  Richmond 

Simalur.  Two  males  and  two  females  measure:  wing,  cf ,  152.5 
(type),  149;  9  ,  144.5,  146;  tail,  tf,  164.5  (type),  169,  9  ,  155,  164.5; 
culmen,  d\  14  (type),  14;  9  ,  13,  14. 

As  well  as  having  certain  differences  in  color  noted  by  Junge  (1936), 
this  race  is  slightly  larger  in  size  than  sumatrana. 

76.  Macropygia  phasianella  hypopercna  Oberholser 

Simalur.  The  type,  an  unsexed  bird  evidently  a  female,  measures: 
wing,  172.5;  tail,  177;  culmen,  17.  This  bird  is  grease  stained  on  the 
upper  back  and  rump.  With  only  one  immature  specimen,  it  is  im- 
possible to  determine  the  validity  of  this  race.  However,  its  resem- 
blance to  the  Nias  race  is  striking. 

77.  Macropygia  phasianella  modiglianii  Salvadori 

Nias.  A  male  and  two  females  measure:  wing,  cf,  184.5,  9,  183, 
185;  tail,  9,  184,  184.5;  culmen,  <?,  17.5,  9,  16,  17. 

This  race  differs  from  emiliana  of  Sumatra  and  Java  principally  by 
larger  size.  There  seems  to  be  a  slight  reduction  also  in  the  amount  of 
the  irregular  barring  on  the  breast. 

78.  Macropygia  phasianella  elassa  Oberholser 

Siberut,  Sipora,  and  North  and  South  Pagi.  The  type,  an  adult 
male  from  North  Pagi,  collected  November  12,  measures:  wing,  179.5; 


RIPLEY:    BIRD    FAUNA    OF   THE    WEST    SUMATRA    ISLANDS  339 

tail,  171  (molt);  culmen,  20.  The  soft  parts  are  marked:  "iris,  pink 
with  an  inner  narrow  blue  circle." 

This  race  is  distinguishable  from  the  available  specimens  of  modi- 
glianii.  In  the  male  the  upper  parts  are  very  similar  but  specimens  of 
elassa  tend  to  have  more  distinct  pale  tips  to  the  feathers  of  the  back, 
scapulars,  and  wing  coverts,  giving  a  slightly  barred  effect.  On  the 
under  surface,  the  iridescent  purple  sheen  of  the  breast  tends  to  be 
confined  to  a  rather  distinct  band,  whereas  in  modiglianii,  the  color  is 
more  generally  dispersed.  The  color  of  the  under  side  of  the  tail 
feathers  seems  to  be  paler,  also,  and  lacking  in  gloss. 

In  the  female,  the  color  differences  are  more  apparent.  The  neck 
and  upper  back  are  distinctly  barred  black  on  buff,  giving  an  effect 
of  more  contrast.  The  tail  seems  to  be  slightly  darker.  On  the  under 
surface  there  is  a  distinct  tendency  to  a  blackish  patch  on  the  throat. 
In  the  male,  this  spot  is  variable  and  sometimes  absent.  In  the  large 
series  of  females  examined,  however,  it  is  always 'present.  The  rest 
of  the  under  surface  seems  to  be  distinctly  darker  than  in  modiglianii. 

79.  Macropygia  phasianella  cinnamomea  Salvadori 

Enggano.  This  race  represents  the  farthest  step  in  a  cline  running 
from  north  to  south  in  the  islands.  The  Nias  population  differs  from 
emiliana  primarily  by  size.  Farther  south  elassa  has  more  distinct 
barring,  a  condition  which  resembles  that  of  immature  birds.  Also 
there  is  a  tendency  towards  reduction  in  the  amount  of  violet  iridescent 
gloss  spread  over  the  plumage  and  the  appearance  of  a  blackish  throat 
patch.  In  cinnamomea  this  condition  is  further  exaggerated.  The  iri- 
descent gloss  has  disappeared.  The  plumage  as  a  whole  appears  faded 
and  washed  out.  The  black  barring  on  the  back  tends  to  break  down. 
The  primaries  are  pale  rufous  on  their  outer  edges  as  in  immature 
specimens  of  the  other  races.  The  black  throat  patch  is  present  as  a 
definite  adult  character. 

An  immature  female  is  uniformly  pale  rufous  brown  all  over,  except 
for  the  upper  back  which  is  irregularly  shaded  with  black,  and  the 
tail  and  inner  margins  of  the  primaries  which  are  blackish. 

Measurements:  wing,  & ,  205.5,  207,  9,  196,  203;  tail,  d\  181.5, 
198,  9  ,  173,  179;  culmen,  d\  22.5,  22.5,  9  ,  20,  20.5. 

80.  Chalcophaps  indica  indica  (Linnaeus) 

A  wide-spread  form  recorded  from  Simalur,  Siumat,  Lasia  (vide 
Chasen,  1935),  Nias,  Tello,  Siberut,  the  Pagi  islands,  and  Enggano. 


340  bulletin:  museum  of  comparative  zoology 

It  will  probably  be  found  to  occur  on  all  the  islands  with  suitable 
forest. 

On  Nias  I  saw  the  Green-wing  Dove  several  times  sunning  and  dust 
bathing  in  the  middle  of  the  small  island  roads. 

81.  Caloenas  nicobarica  nicobarica  (Linnaeus) 

Recorded  from  Kokos,  Simalur,  Lasia,  Babi,  Nias,  Batu  Ids.,  and 
Mirabau,  an  islet  off  Enggano. 

This  is  primarily  a  bird  of  the  small  coral  islands  off  shore,  but 
Nieuwenhuisen  and  von  Rosenberg  record  it  as  common  in  the 
Lagoendi  Bay  region  of  south  Nias,  near  the  present-day  Telokdalem. 


Family  PSITTACIDAE,  Parrots 

Seven  psittacine  forms  occur  on  the  west  Sumatra  islands  of  which 
six  are  endemic.  The  species  Psittacula  alexandri  has  no  representa- 
tives on  Sumatra.  The  three  races  found  on  these  islands  are  closest 
to  the  population  of  the  Andaman  Islands.  Like  that  race  they  differ 
from  the  bird  of  the  mainland  primarily  by  larger  size  as  well  as  certain 
color  characters. 

The  species  Psittacula  longicauda  found  on  Sumatra  as  well  as  the 
Andamans  and  Nicobars,  has  a  single  highly  specialized  representative 
on  these  islands  on  Enggano.  It  differs  by  much  larger  size  and  rather 
juvenal  color  characters. 

There  are  two  forms  of  Psittinus  cyanurus  on  these  islands.  One, 
from  Simalur  is  not  only  much  larger  than  the  Sumatran  bird,  but 
has  certain  striking  color  differences.  The  other,  from  the  Mentawi 
and  Pagi  Islands  differs  from  the  Sumatra  bird  only  in  larger  size. 


82.  Psittacula  alexandri  cala  (Oberholser) 

Simalur.  The  type,  an  adult  male  taken  October  21,  measures: 
wing  185,  tail  198.5.  The  rest  of  the  series  collected  by  Abbott  in 
October  and  November  measure:  wing,  d\  173.5  (worn)-177.5,  9, 
168.5-177.5;  tail,  cf,  179.5-196,  9,  170-182. 

This  race  resembles  abbotti  from  the  Andaman  islands  very  closely. 
The  only  tenable  difference  seems  to  be  that  the  pinkish  color  of  the 
breast,  particularly  in  the  females,  is  more  pure,  less  suffused  with 
lilac.  Other  differences  can  apparently  be  matched  in  a  large  series. 


RIPLEY:    BIRD    FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  341 

From  fasciata  this  race  differs  by  larger  size,  lack  of  any  trace  of 
green  on  the  forehead,  and  the  less  intense  shade  of  green  on  the  back. 

This  species  is  abundant  in  the  west  Sumatra  islands  nocking  to- 
gether and  nesting  throughout  the  year. 

83.  Psittacula  alexandri  major  (Richmond) 

Lasia  and  Babi.  The  type,  an  adult  male  from  Babi,  January  14, 
measures:  wing,  196.5,  tail,  230.5.  Other  birds  from  the  two  islands 
measure:  wing,  d\  189.5,  196.5,  201.5,  9,  186,  188.5;  tail,  d\  218.5, 
229,  9  ,  196,  200.5. 

This  is  a  very  large  race  differing  from  cala  primarily  in  size, 
although  the  males  seem  to  show  a  reduction  in  the  light  bluish  suf- 
fusion on  the  lower  abdomen  and  vent.  One  female  (U.S.N.M. 
179112)  has  a  thin  ring  of  pink  extending  from  the  sides  of  the  neck 
dorsally  around  the  nape  between  the  bluish-gray  crown  and  the 
green  back. 

Soft  parts:  iris,  pale  yellow  with  an  inner  dull  green  ring;  upper 
mandible  (cf),  red;  cere,  greenish  leaden,  dull  greenish;  feet,  pale 
green,  pale  dusty  green. 

84.  Psittacula  alexandri  perionca  (Oberholser) 

Nias.  The  type,  an  adult  male  collected  February  22,  measures: 
wing,  194.5;  tail,  211.5.  Another  male  and  two  females  measure :  wing, 
tf,  182,  9  ,  177,  183.5;  tail,  <?,  189.5,  9  ,  (molt),  170. 

This  race  occupies  a  somewhat  intermediate  position  as  regards  cala 
and  major  in  size.  Like  major  the  males  have  less  of  the  heavy  bluish 
suffusion  on  the  lower  abdomen  and  vent.  The  color  of  this  area  is 
somewhat  brighter  than  in  major. 

In  June  these  birds  were  nesting,  using  holes  made  high  up  in  the 
trunks  of  dead  coconut  palms. 

85.  Psittacula  longicauda  modesta  (Fraser) 

Enggano.  The  two  species,  alexandri  and  longicauda,  have  an  extra- 
ordinary distribution  in  the  greater  Sunda  area.  It  is  particularly 
interesting  that  alexandri,  a  species  found  away  from  the  mainland, 
principally  on  small  islands,  should  have  reached  Java.  This  argues 
a  long  history  for  the  range  of  the  species.  And  yet  it  is  a  representa- 
tive of  longicauda  which  has  reached  Enggano,  the  most  isolated  of 
the  west  Sumatra  islands. 


342 


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The  race  modesta. occupies  one  end  of  a  cline  starting  with  longicauda 
of  the  mainland,  Sumatra,  and  Borneo.  The  cline  consists  principally 
of  color  characters  which  may  be  summarized  as  follows: 


Subspecies 

Loral 
stripe 

Crown 

Nape 

Back  vs. 
Scapulars 

Size 
Wing  d"  cf 

longicauda 

indicated 

emerald 
green 

pink,  like 
cheeks 

t 

strong 
contrast 

small 

defontainei 

indicated 

yellowish 

emerald 

green 

paler 

strong 
contrast 

larger 

tytleri 

present 

yellowish 
green 

lacking 

weaker 
contrast 

larger 

nicobarica 

present 

chromium 
green 

lacking 

weak 

contrast 

larger 

modesta 

extending 

over 

nostrils 

indistinctly 
green 

present 

very 

pale 

weakest 
contrast 

largest 
199.5-209 

The  series  collected  by  ^Abbott  measures:  wing,  cf,  199.5-209 
(205.6),  9,  195.5-209.5  (201.1);  tail,  d71,  182  (worn)-233,  9,  131- 
151.5 

Molting  specimens  were  collected  in  October  and  November  and  a 
fully  grown  immature  female  November  11.  The  soft  parts  are  marked 
as:  iris,  yellow,  inner  circle  green;  upper  mandible,  red,  lower,  horn 
brown,  black  (cf);  feet,  greenish  leaden,  slaty-green,  greenish. 

A  male  and  a  female  are  heavily  stained  on  the  under  parts,  pre- 
sumably from  eating  fruit. 

86.  Psittinus  ctanurus  abbotti  Richmond 

Simalur  and  Siumat.  This  very  distinct  form  lies  somewhere  on 
the  borderline  between  a  species  and  a  subspecies.  Primarily  it  reflects 
the  speciation  trend  of  these  islands  by  being  much  larger  than  cya- 
nurus.  Also,  however,  there  are  important  color  characters  which  set 
it  off  from  the  typical  race. 

Adult  male:  the  crown  is  a  more  uniform  blue  than  in  cyanurus, 
lacking  the  gray  wash  on  the  nape.  Also  there  is  a  sprinklingof  greenish 


RIPLEY:   BIRD    FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  343 

iridescent  feathers  on  the  forehead  and  the  ocular  area.  The  grayish 
black  back  and  scapulars  of  cyanurus  are  lacking,  being  replaced  with 
green.  The  mid  portion  of  the  back  is  bright  blue  instead  of  pale  dull 
purplish  blue  and  the  rump  and  upper  tail  coverts  are  green  instead  of 
blue.  On  the  under  surface  greenish  yellow  replaces  the  yellowish  gray 
breast,  abdomen,  and  vent,  and  the  blue  thighs  of  cyanurus. 

Adult  female:  above  this  bird  is  almost  uniformly  green,  lacking 
the  dull  brown  head  of  the  typical  race.  Below,  also,  abbotti  is  uni- 
formly greenish  yellow. 

The  type,  an  adult  male  taken  in  December,  measures:  wing,  145; 
tail,  58.5;  culmen  (from  cere)  22.5;  tarsus,  13.  Iris,  pale  yellow;  cere, 
dull  green;  feet,  greenish. 

A  series  of  males  and  females  measure:  wing,  c? ,  139-147.5  (143.5); 
cf,  im.,  139;  9  ,  134.5-141  (137.6). 

An  immature  male  collected  in  October  has  an  indistinct  tinge  of 
blue  on  the  forehead.  An  adult  male  (U.S.N.M.  179109)  has  a  spot  of 
dull  red  similar  to  that  of  the  humeral  patch  among  the  blue  feathers 
of  the  nape. 

This  is  a  rather  tame  little  parrot,  common  locally  when  the  wild 
fig  trees  are  fruiting. 

87.  Psittinus  cyanurus  pontius  Oberholser 

Siberut,  Sipora,  and  South  Pagi.  This  race  is  distinguished  from 
cyanurus  by  larger  size.  In  a  large  series,  any  color  differences  are 
shown  to  vary  within  the  species.  A  male  from  Siberut  (U.S.N.M. 
279754)  has  a  very  dark  blackish  back.  Soft  parts:  "iris,  pale  yellow; 
upper  mandible,  red  tipped  with  yellow,  lower,  olive  horn;  cere,  dull 
leaden  olive;  feet,  olive." 

The  type,  an  adult  male  collected  in  December,  measures:  wing, 
127;  tail,  50.5;  culmen,  20.75.  The  rest  of  the  series  measures:  wing, 
<? ,  127-134.5  (131.5);  9  ,  129-132;  tail,  cf ,  50.5-54.  Birds  from  Su- 
matra and  the  Malay  Peninsula  measure:  wing,  cf ,  117-125  (121.6), 
9,  115-123;  tail,  44-47. 

88.  Loriculus  galgulus  galgulus  (Linnaeus) 
Synonym:  Loriculus  galgulus  lamprochlorus  Oberholser 
Synonym:  Loriculus  galgulus  dolichopterus  Oberholser 

Tuangku,  Nias,  Pini,  Tello,  Siberut,  Sipora,  Enggano. 

The  type  of  lamprochlorus,  an  adult  male  collected  on  Nias,  March 


344  bulletin:  museum  of  comparative  zoology 

14,  measures:  wing,  79;  tail,  32.5;  culmen  (from  cere),  10.5.  This  race 
was  described  by  Oberholser  (1912)  as  being  "decidedly  smaller,  and 
the  colors  averaging  paler;  female  with  green  color  more  yellowish". 
A  series  of  birds  from  Sumatra,  Borneo,  and  Java  (three  specimens 
which  may  well  be  cage  birds)  measure:  wing,  cT,  78.5-83  (80.3). 
The  color  characters  also  disappear  in  a  series. 

The  type  of  dolichopterus,  a  female  from  Enggano  collected  Novem- 
ber 6,  measures:  wing,  83.5;  tail,  34.5;  culmen  (from  cere),  11.5.  A 
series  of  females  from  Sumatra,  Borneo,  and  Java  (see  anted)  measure: 
wing,  77-84.5  (81.4).  This  race  was  described  as  being  larger  and 
darker,  but  these  differences  do  not  appear  in  comparison  with  a 
large  series  of  birds.  Junge  (1938)  reached  the  same  conclusion. 

The  wings  of  some  of  these  specimens  have  not  been  fully  pulled 
back  after  skinning.  In  the  case  of  the  type  of  dolichopterus  this  may 
have  accounted  for  the  impression  that  it  was  a  larger  bird. 

Family  CUCULIDAE,  Cuckoos 

Fifteen  species  have  been  recorded  from  these  islands  of  which  two 
are  migrants.  Of  the  remaining  species  four  are  endemic  on  the  islands. 
Of  these  four  races,  two  differ  by  color  alone  while  the  other  two  differ 
only  by  larger  size.  All  the  races  are  closely  related  to  representatives 
of  the  species  found  on  Sumatra. 

89.  Cuculus  fugax  fugax  Horsfield 

Pini,  Siberut.  An  immature  specimen  from  Siberut  collected 
October  first,  apparently  belongs  to  this  race.  Chasen's  record  for 
nisicolor  (1935)  may  well  be  founded  on  this  and  another  immature 
bird  taken  by  Kloss.  The  bird  measures:  wing,  167.5;  bill  (from  gape), 
26.5.  Some  of  the  slaty  adult  feathers  are  beginning  to  appear  on  the 
forehead,  cheeks,  and  nape.  There  are  several  white  feathers  also  on 
the  nape. 

90.  Cuculus  micropterus  concretus  S.  Muller 

Tana  Bala.  An  adult  male  collected  February  11,  tail  lacking,  is 
the  first  record  of  this  species  from  the  west  Sumatra  islands.  It  meas- 
ures: wing,  191.5;  culmen,  22.5.  The  soft  parts  are  recorded  as:  "iris, 
ochreous,  rim  of  eyelid  yellow;  lower  mandible,  gray;  gape,  yellow; 
feet,  yellow."  Compared  with  a  male  from  Trang  and  a  female  frojn 
east  Sumatra  collected  in  January  and  February,  this  specimen  is  dis- 


RIPLEY:    BIRD    FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  345 

tinctly  more  gray  on  the  back,  wing  coverts,  rump,  and  upper  tail 
coverts.  The  underparts  seem  to  be  identical.  It  is  unfortunate  that 
no  more  specimens  were  secured. 

91.    CUCULUS  SATURATUS  SATURATUS  Blyth 

A  migrant  from  southern  Asia  discussed  by  Junge  (1931)  in  a  careful 
revision  of  the  species.  He  lists  two  specimens  of  this  race  from 
Simalur  taken  in  February.  The  other  record  for  the  west  Sumatra 
islands  is  Nias  (Biittikofer  1896,  p.  171). 

92.  Penthoceryx  sonneratii  fasciolatus  (S.  M  tiller) 

Pini,  Tana  Massa.  A  male  and  a  female  collected  by  Kannegieter 
in  September  and  October  on  these  two  islands  measure:  wing,  d* 
(worn)  114,  9  111.  These  birds  are  indistinguishable  from  two  males 
from  Trang,  a  locality  certainly  within  the  range  of  malayanus  Chasen 
and  Kloss. 

93.  Cacomantis  merulinus  threnodes  Cabanis  and  Heine 
Synonym:  Cacomantis  merulinus  subpallidus  Oberholser 

Nias,  Siberut,  and  Enggano.  The  type  of  subpallidus,  an  immature 
male  from  Nias,  is  a  rather  pale  bird,  but  an  adult  male  from  Nias  is 
similar  to  a  male  from  Borneo.  I  believe  that  this  slight  difference  in 
coloration  is  due  to  individual  variation.  Two  adult  males  from  Nias 
and  Siberut  measure:  wing,  101,  101.5. 

The  characteristic  ascending  notes  of  the  brain-fever  bird  are  often 
heard  on  Nias  in  open  areas  of  garden  and  cultivation. 

94.  Cacomantis  variolosus  sepulcralis  (S.  Mtiller) 

Simalur,  Enggano.  A  male  molting  into  adult  plumage  and  an  im- 
mature female  were  taken  by  Abbott  on  Simalur,  December  14  and 
November  30.  The  soft  parts  were  noted  as:' "iris,  d*  gray  brown;  9 
reddish  brown  becoming  gray  externally;  eyelids,  d1  greenish  yellow; 
bill  black,  brownish  yellow  at  base  of  lower  mandible;  inside  of  mouth 
orange;  feet  yellow." 

95.  Chalcites  xanthorhynchus  xanthorhynchus  (Horsfield) 
Only  recorded  from  Simalur  and  Sipora. 


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96.  SURNICULUS  LUGUBRIS  DICRUROIDES  (Hodgson) 

A  female  collected  on  Nias  March  15  evidently  belongs  to  this  large 
raigatory  race.  It  measures :  wing  141.5,  tail  135,  culmen  22.  The  soft 
parts  are  labelled:  "iris  dark  brown,  bill  black,  feet  dark  leaden." 

97.  Surniculus  lugubris  barussarum  Oberholser 

Tana  Bala.  The  type  of  barussarum,  an  adult  female  taken  on  Tana 
Bala  in  February,  measures:  wing  136,  tail  127,  culmen  22.  Another 
female  taken  in  the  same  month  measures:  wing  137,  tail  (worn)  120, 
culmen  22.  Both  birds  have  distinctly  forked  tails,  the  lateral  feathers 
in  the  type  being  6mm.  longer  than  the  central  ones. 

This  race  was  named  (1912,  p.  5.)  as  being  smaller  with  the  bill 
relatively  larger  than  I.  lugubris.  Actually  two  males  of  lugxibris  from 
east  Java  have  the  following  measurements:  wing  124,  127,  culmen 
20.5,  21.  Robinson  (Birds  Malay  Penin.  /,  1927,  p.  140.)  gives  meas- 
urements for  brachyrus  of  the  Malay  Peninsula,  Borneo,  and  Sumatra 
which  are  very  close  to  these  two  birds  from  Tana  Bala.  A  female 
from  Singkep  Island,  east  Sumatra  taken  in  May  measures  only: 
wing  128,  tail  123,  culmen  20.5.  If  this  specimen  may  be  taken  as  an 
example  of  typical  brachyurus,  then  I  presume  that  banissarum  repre- 
sents a  large  island  form.  Larger  series  may  show,  however,  that  these 
measurements  are  not  distinctive,  in  which  c&sebarussarum  must  stand 
as  the  name  for  the  birds  from  the  southern  Malay  Peninsula,  Borrfeo, 
Sumatra,  and  the  Batu  Ids. 

98.    EUDYNAMIS  SCOLOPACEA  SIMALURENSIS  Junge 

Kokos,  Simalur,  Lasia,  Babi.  Junge  (1936)  characterizes  this  race 
as  differing  from  malayana  by,  "smaller  wing  and  in  the  9  also  by 
the  darker  coloured  and  bigger  spots  on  the  upper  side  and  the  more 
strongly  red-brown  washed  under  parts". 

As  to  size  I  cannot  entirely  agree  with  this  diagnosis.  A  male  from 
Babi  measures:  wing  211,  tail  197,  culmen  36,  while  a  male  from 
Simalur  in  the  collection  of  the  Museum  of  Comparative  Zoology  has 
a  wing  measurement  of  only  195  according  to  Mr.  Peters.  A  series  of 
males  of  malayana  measure:  wing  203-212.5.  However,  a  single  pos- 
sibly immature  female  from  Simalur  is  distinctly  more  rufous  above 
and  below  than  any  female  of  malayana.  In  this  respect  it  is  very 
close  to  females  of  mindanensis.   Fortunately  the  bill  of  mindanensis 


RIPLEY:   BIRD    FAUNA   OF  THE   WEST    SUMATRA    ISLANDS  347 

is  noticeably  smaller.   The  record  for  Lasia  is  a  sight  one,  (Richmond, 
1903). 

99.  Eudynamis  scolopacea  malayana  Cabanis  and  Heine 

Nias,  Tello,  Tana  Massa,  Pagi  islands.  Two  males,  an  immature 
male  and  three  females  from  Tello  and  Tana  Massa  measure:  wing, 
c?  217.5,  218;  9  im.  213,  9  209,  212,  213. 

There  is  a  good  deal  of  individual  variation  in  size,  bill  dimensions, 
and  color  in  this  species.  These  birds  are  large  but  not  significantly 
so.  The  soft  parts  are  recorded  as:  iris  red;  bill  gray,  greenish  gray; 
feet  gray. 

The  record  for  the  Pagi  islands  is  not  founded  on  any  specimen  so 
far  as  I  know,  but  undoubtedly  these  birds  could  be  found  on  almost 
any  of  the  small  coral  islands  of  the  area,  especially  in  the  breeding 
season. 

100.  Rhopodytes  sumatranus  rodolphi  Ripley 

Pini.  The  type,  an  adult  male  collected  October-November  1896, 
measures :  wing  154,  tail  237,  culmen  37.5.  The  soft  parts  are  recorded 
as:  iris  light  blue,  bill  yellowish  green,  feet  gray. 

Although  only  a  single  specimen  has  been  taken  on  the  west  Sumatra 
islands,  its  bill  measurements  are  significantly  larger  than  a  series  of 
specimens  from  Sumatra  or  the  Malay  Peninsula  and  adjacent  islands. 
In  color  there  seems  to  be  no  perceptible  differences  between  this  and 
Sumatra  or  southern  Malay  Peninsula  birds.  This  specimen  has  very 
little  rufous  on  the  abdomen,  but  I  believe  that  individual  variation 
or  skinning  will  account  for  it. 

101.  Rhinortha  chlorophaea  facta  Ripley 

Tana  Massa.   The  type,  an  adult  male  collected  February  20,  1903, 
measures:  wing  123.25,  tail  182,  culmen  29,  tarsus  28.5.    A  female 
(A.N.S.P.  56256)  collected  August  24,  1896,  measures:  wing  124,  tail 
179.5,  culmen  30.5,  tarsus  28.5.   The  soft  parts  are  given  as:  "iris,  9 
blue  gray;  bill,  cf  dark  greengage  green,  9  ,  green;  feet,  c?  slaty,  9 
gray." 

These  birds  are  noticeably  larger  than  any  other  specimens  of  the 
species  in  the  Museum's  series.  Except  for  Banguey  I.  off  the  coast 
of  North  Borneo  this  bird  has  never  been  reported  on  any  of  the  small 
islands  in  the  Greater  Sunda  area. 


348  bulletin:  museum  of  comparative  zoology 

102.  Rhamphococcyx  curvirostris  oenicaudus 
(Verreaux  and  Verreaux) 

Siberut,  Sipora,  North  and  South  Pagi.  This  race  differs  from 
erythrognathus  of  the  Malay  Peninsula  and  Sumatra  by  having  the 
greenish  iridescence  on  the  upper  surface  rather  darker  and  more 
bronzy,  extending  up  onto  the  nape  and  crown  and  serving  to  darken 
the  gray  of  those  areas.  The  tail  lacks  the  terminal  brown  patch 
characteristic  of  the  other  members  of  the  species. 

Below  the  greenish  iridescence  extends  up  over  the  abdomen.  The 
under  surface  of  the  tail  is  greenish  blue  not  brown.  There  seems  to 
be  no  size  difference  within  the  species. 

Five  males,  seven  females,  and  three  sex  unidentified,  measure: 
wing,  cf ,  166-181,  9  165-178,  o  171-179.  Soft  parts  are  recorded  as: 
"iris,  c?  bluish  white,  pale  blue,  pale  china  blue,  9  dark;  bill,  maxilla 
apple  green;  nostril  black,  pale  green;  mandible  green,  base  dull 
crimson,  dark  red;  orbital  skin  scarlet,  red;  feet  plumbeous." 

103.  Centropus  sinensis  bubutus  Horsfield 

Nias,  Siberut.  Four  males  and  one  unsexed  from  Nias  collected  in 
February  and  March  measure:  wing,  &  212,  215,  218,  221,  o  220,  tail 
<?  272,  278,  278,  o  285.  The  variation  in  these  birds  from  purplish 
to  greenish  iridescence  about  the  head,  neck,  and  breast  varies  with 
the  freshness  of  the  plumage. 

Family  TYTONIDAE,  Barn  Owls 

The  only  species  of  this  family  found  in  the  area  is  the  small  Bay 
Owl  of  the  greater  Sunda  Islands  which  is  not  uncommon  on  Nias. 

104.  Phodilus  badius  badius  (Horsfield) 
Nias.   This  species  has  only  been  recorded  from  Nias  where  it  is 
not  uncommon.   For  further  comments  and  field  notes  see  de  Schau- 
ensee  and  Ripley  (1939,  p.  402.). 

Family  STRIGIDAE,  Owls 

Six  forms  of  owls  are  found  on  the  west  Sumatra  islands,  all  of  which 
are  apparently  endemic.  Two  differ  from  their  congeners  in  Sumatra 
and  Borneo,  primarily  by  color  differences.    Three  of  the  remaining 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  349 

forms  are  distinctly  larger  than  their  nearest  relatives  on  the  Malay 
Peninsula  or  Sumatra,  as  well  as  differing  slightly  in  color.  One  race 
is  smaller  than  its  Sumatran  congener. 

105.  Otus  scops  umbra  (Richmond) 

Simalur.  The  type  and  unique  specimen,  an  adult  male,  was  col- 
lected by  Abbott  November  29,  1901.  It  measures:  wing  143,  tail  61, 
culmen  20,  tarsus  24.  The  soft  parts  are  recorded  as:  iris  greenish 
yellow;  bill  pale  brown,  black  at  tip;  feet  pale  fleshy  brown.  The  wing 
formula  is  4>3>5>2>6>7>1. 

This  bird  is  presumably  in  the  rufescent  phase.  The  outer  scapulars 
have  conspicuous  white  spots  and  the  feathers  of  the  lower  abdomen, 
vent,  and  flanks  are  freckled  and  mottled  with  white, russet,  and  black 
in  a  way  that  seems  characteristic  of  the  scops  group. 

The  stomach  contained  insects. 

106.  Otus  scops  enganensis  Riley 

Enggano.  A  single  female,  the  type,  was  collected  by  Abbott,  No- 
vember 12,  1904.  Its  measurements  are:  wing  (worn),  tail  73,  culmen 
23,  tarsus  30.  The  tips  of  the  primaries  are  completely  worn  off,  a 
condition  that  was  not  mentioned  in  the  original  description.  The 
wings  of  five  birds  collected  by  deJong  (Junge  1938)  measure:  c? , 
160,  163;  9  ,  163,  163,  166.  The  soft  parts  in  the  type  are  recorded  as : 
"iris  greenish  yellow,  feet  pale  brownish  fleshy." 

The  type  is  a  bird  in  the  rufescent  phase  of  plumage.  In  his  dis- 
cussion of  this  form,  Junge  (I.e.)  showed  that  the  brownish  or  gray- 
brownish  individuals  are  very  close  to  malayanus  and  sunia.  The 
under  parts,  however,  are  more  uniformly  dark  rufous  than  any 
rufescent  individual  of  scops  I  have  seen,  except  for  the  type  of  umbra, 
which  is  the  most  uniformly  dark  of  all. 

107.  Otus  bakkamoena  mentawi  Chasen  and  Kloss 

Siberut,  Sipora,  North  Pagi.  In  their  original  description  (1926), 
Chasen  and  Kloss  neglected  to  state  that  this  race  differed  from  true 
lempiji  by  much  larger  size.  A  series  of  these  birds  measure:  wing, 
c?  157>  165,  9  165,  165.5,  9  im.  160;  tail,  tf  77.5,  82,  9  72,  80,  80, 
9  im.  76.  Ten  specimens  of  lempiji  from  Java,  Sumatra,  and  the 
Malay  Peninsula  measure:  wing,  cf  141-150.5  (145.2),  9  140-151.5 
(145.1);  tail,  d*  65-70  (67.2),  9  65-67.5  (66.3).. 


350  bulletin:  museum  of  comparative  zoology 

The  soft  parts  of  these  birds  are  listed  as:  "iris,  brown  (3),  yellow 
(2),  feet  gray."  The  color  differences  noted  in  the  original  description 
are  very  noticeable  in  this  series. 

108.  Ketupa  ketupu  minor  Biittikofer 

Nias.  Compared  with  ketupu  from  adjacent  areas,  three  males  and 
a  female  from  Nias  show  the  following  measurements: 

wing  tail 

Nias  cf  304-330.5  (316.1)  143-164  (153.5) 

9                                     304  147 

Sumatra1  cf  320-349  (338.6)  158-171  (165.4) 

9                          335,  338  158 

Borneo  c?  312-343  (330.1)  153-164.5  (158.4) 

9                                   —  — 

Java  &  341-357  (348.4)  157-169  (164) 

9                               358,  362  170,  176 

From  the  above  it  will  be  seen  that  minor  is  a  tenable  race  but  that 
there  is  a  good  deal  of  overlap  with  Sumatra  and  Borneo  birds.  Two 
of  the  Nias  males  are  darker  on  the  breast  than  any  specimen  of  our 
series  of  ketupu. 

Three  young  were  taken  in  late  March.  The  plumage  is  rather  uni- 
form dark  buffy  all  over  except  for  the  thighs  which  are  whitish.  Both 
upper  and  under  parts  are  marked  with  the  narrow  dark  brown  shaft 
streaks  found  in  the  adult  only  on  the  under  parts. 

109.  Strix  leptogrammica  nyctiphasma  Oberholser 

Bangkaru.  The  type,  an  adult  male  collected  in  January,  measures: 
wing  299.5,  tail  154.  A  female  taken  at  the  same  time  measures:  wing 
307,  tail  167. 

The  type  differs  from  a  male  from  the  Kapuas  River,  west  Borneo 
(w.  298,  t.  168)  by  having  the  cheeks  somewhat  paler  and  the  dark 
barring  on  the  underparts  slightly  heavier,  the  bars  being  closer  to- 
gether. In  size  there  is  no  difference  and  the  color  differences  are  so 
slight  that  a  larger  series  may  well  show  these  forms  to  be  identical. 

From  myrtha  of  Sumatra  these  birds  differ  primarily  by  smaller 
size.  This  bird  will  probably  be  found  one  day  on  Babi  and  Simalur, 
provided  there  is  some  original  forest  left  for  them. 

1  have  padded  out  my  series  with  measurements  taken  in  the  same  manner  by  Mayr  (Bull. 
Raffles  Mus.,  No  14,  1938,  p.  13)  and  de  Schauensee  and  Ripley  (Proc.  Acad.  Sci.  Philadelphia. 
91,  1939,  p.  324.) 


RIPLEY:    BIRD    FAUNA    OF   THE   WEST   SUMATRA   ISLANDS  351 

110.  Strix  leptogrammica  niasensis  Salvadori 

Xias.  A  male  and  female  (M.C.Z.  194794,  194795)  collected  in 
July  1937  by  Barbara  Lawrence  measure:  wing,  a71  273,  9  280.5; 
tail,  d"  151.5,  9  156.5.  The  male  is  an  immature  probably  completing 
the  post  juvenal  molt  {vide  Robinson,  Birds  Malay  Benin.,  2,  1928, 
p.  36).  The  feathers  of  the  posterior  part  of  the  crown  and  nape  and 
a  few  feathers  in  a  ring  on  the  neck  below  are  pale  whitish  buff  in  color 
and  fluffy. 

The  iris  is  recorded  as  brown  by  Biittikofer,  chestnut  by  Salvadori 
(1887).  Salvadori  (I.e.)  lists  the  bill  and  feet  as  pearl  gray  or  sky  blue. 


Family  CAPRIMULGIDAE,  Nightjars 

There  are  records  of  three  forms  from  these  islands  of  which  one 
has  been  separated  on  color  characters,  and  one,  Eurostopodus  tem- 
minckii,  shows  an  unconfirmed  tendency  towards  larger  size.  The 
endemic  form,  Eurostopodus  viaCrotis  jacobsoni  has  no  close  geograph- 
ical relatives  except  cerviniceps  from  the  Malay  Feninsula. 

111.  Eurostopodus  temminckii  (Gould) 

Nias.  Two  males  taken  by  Abbott  in  March  measure:  wing  211, 
213;  tail  133.5,  135.  Two  males  from  Johore  and  Banka  measure: 
wing  201,  203;  tail  117,  126,  while  three  males  from  north  Sumatra 
are  recorded  by  de  Schauensee  and  myself  (I.e.,  1939)  as  having 
wings  of  197,  198.5,  and  201.5  mm.  However,  Robinson  and  Kloss 
(Journ.  Fed.  Malay  States  Mus.,  11,  1924,  p.  242)  give  measurements 
for  males  up  to  218. 

In  these  two  specimens  the  buffy  whitish  subterminal  bars  forming  a 
ring  on  the  nape  are  narrower  than  in  any  other  specimens  in  the 
National  Museum  series.  Also  one  specimen  (180824)  has  very  large 
black  spots  on  the  crown. 

112.  Eurostopodus  macrotis  jacobsoni  (Junge) 

Simalur.  According  to  the  description  (1936)  a  darker  bird  than 
macrotis  from  the  Bhilippines  or  cerviniceps  of  the  Malay  Beninsula. 
It  is  most  interesting  to  find  this  population  so  isolated  from  the  rest 
of  the  species.  Further  collecting  might  reveal  that  it  occurs  in  the 
Andamans  or  Nicobars. 


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113.  Caprimulgus  affinis  affinis  Horsfield 

Recorded  from  Nias  by  Nieuwenhuisen  and  von  Rosenberg  (1863) 
under  the  name  C.  maculatus. 


Family  MICROPODIDAE,  Swifts      . 

Six  races  of  swifts  are  found  on  the  west  Sumatra  islands  of  which 
two  are  endemic.  One  race  differs  in  color  alone,  and  the  second  differs 
in  color  and  smaller  size.  Both  races  are  closely  related  to  a  wide- 
spread form  found  throughout  the  greater  Sunda  area. 

114.  Collocalia  lowi  lowi  (Sharpe) 

Simalur,  Babi  (sight),  Nias.  A  male  from  Simalur  taken  May  5  is 
listed  by  Junge  with  a  wing  measurement  of  131. 

115.  Collocalia  fuciphaga  fuciphaga  (Thunberg) 

Nias.  A  single  female  taken  in  June  is  the  only  record  of  this  species 
for  the  west  Sumatra  islands. 

116.  Collocalia  vestita  vestita  (Lesson) 
Synonym:  Collocalia  fuciphaga  aerophila  Oberholser 

Simalur,  Nias,  Sipora.  The  type  of  aerophila,  an  adult  male  from 
Nias,  collected  March  16,  measures:  wing  115;  tail,  shortest  rectrix 
48,  longest  rectrix  53.  A  male  and  a  female  from  Simalur  and  a  female 
from  Sipora  measure:  wing,  d71  112.5,  9  116.5,  116;  tail,  <?  46  and 
50,  9  41.5  and  47,  49  (incomplete).  Compared  with  a  male  from 
Johore,  I  can  see  no  significant  differences  in  these  specimens. 

117.  Collocalia  esculenta  cyanoptila  Oberholser 

Simalur.  Junge  (1936)  records  three  specimens  taken  on  Simalur 
in  April  and  July  as  belonging  to  this  race.  He  gives  their  measure- 
ments as  follows :  wing,  cf  107,  109,  9  102;  tail,  cf  41,  41.  The  type 
and  another  female  of  cyanoptila  from  Bunguran-L,  Natunas,  measure: 
wing,  9  103.5,  103.5;  tail,  41.5,  39.5  (molt).  Both  birds  are  in  rather 
worn  plumage  which  accounts  for  the  admixture  of  purple  and  greenish 
gloss  mentioned  by  Oberholser  in  his  original  description  (Proc.  Acad. 
Nat.  Sci.  Phila.,  58,  1906,  p.  205). 


RIPLEY:    BIRD    FAUNA    OF   THE   WEST    SUMATRA   ISLANDS  353 

118.  Collocalia  esculenta  vanderbilti  de  Schauensee  and  Ripley 

Nias.  The  male  type  and  a  female  measure :  wing,  cf  99,  9  97;  tail, 
cf  39,  9  37.  This  race  is  smaller  than  oberholseri  and  somewhat  bluer 
above.  These  little  swiftlets  are  common  over  jungle  trails  and  roads, 
especially  in  the  early  morning  and  evening. 

119.  Collocalia  esculenta  oberholseri  Stresemann 

Tana  Massa,  Sipora  and  North  Pagi.  The  type,  an  adult  male  from 
North  Pagi,  taken  in  November,  measures:  wing  105.5,  tail  42.5. 
Other  birds  from  Sipora  and  North  Pagi  measure:  wing,  cf  105; 
9  98,  105;  o  105,  105.5;  tail,  cf  42.5;  9  40.5,  41.5;  o  42,  42.5. 

From  cyanoptila  this  race  differs  by  being  more  greenish  above  and 
with  the  breast  and  upper  abdomen  paler  below  with  more  tips  among 
the  brownish  gray  of  that  area. 


Family  HEMIPROCNIDAE,  Tree  Swifts 

Two  races  of  Tree  Swifts  are  found  on  these  islands  of  which  one  is 
endemic.  It  differs  from  its  large  island  and  mainland  relatives  by 
larger  size. 

120.  Hemiprocne  longipennis  perlonga  (Richmond) 
Synonym:  Hemiprocne  longipennis  ocyptera  Oberholser 
Synonym:  Hemiprocne  longipennis  thoa  Oberholser 

Simalur,  Nias,  Pini,  Tana  Massa,  South  Pagi,  Enggano.  The  type 
of  perlonga,  an  adult  female,  was  collected  on  Simalur  January  2, 
1902.  It  measures:  wing  183.5,  tail  112.5.  The  type  of  ocyptera,  an 
adult  male  collected  March  23,  1905  on  Nias  measures:  wing  169. 
The  type  of  thoa,  an  adult  male  from  Pini  taken  March  7,  1903, 
measures:  wing  177.5,  tail  102.  These  last  two  races  were  described  in 
a  lucid  fashion  by  Oberholser  (1912,  pp.  7,  8)  without  being  compared 
to  each  other,  although  they  came  from  closely  neighboring  islands. 

A  total  of  seventeen  specimens  from  the  west  Sumatra  islands  have 
wing  measurements  as  follows:  cf  164-180  (173.1),  9  169.5-183.5 
(174.9).  Of  this  series  the  birds  from  Nias  are  the  smallest:  cf  164- 
176  (169),  9  169.5.  This  is  probably  due  to  the  fact  that  Nias  of  all 
the  islands  of  the  group  seems  to  be  the  most  closely  connected  to 
Sumatra  from  a  faunistic  point  of  view.   Whether  in  the  case  of  Hemi- 


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procne  this  might  be  due  to  swamping  is  certainly  a  possibility  in  the 
case  of  this  distinctly  aerial  species. 

I  have  examined  forty-one  specimens  of  longipennis  from  the  Malay 
Peninsula,  Rhio  and  Lingga  islands,  Sumatra,  Natunas,  Anambas, 
Borneo,  Banka,  and  Java.  Seventeen  males  measure:  159-170.5 
(164.3).  Exclusive  of  molting  birds,  twenty-three  females  measure: 
155-173.5  (166).  From  these  figures  it  will  be  seen  that  perlonga  is  a 
tenable  subspecies.  However,  it  seems  to  me  that  in  general  this 
species  has  been  split  into  far  too  many  subspecies.  The  type  of 
anochroa  Oberholser,  an  adult  female  from  the  Natunas,  has  a  greenish 
suffusion  in  the  chest  region  and  in  general  is  somewhat  darker  than 
average  on  the  underparts.  But  in  this  it  can  be  matched  by  birds 
from  the  Malay  Peninsula,  Sumatra,  and  Borneo. 

Stresemann's  original  description  of  the  race  harterti  (Novit.  Zool., 
20,  1913,  p.  339)  indicates  that  he  did  not  have  any  Bornean  material, 
for  he  gives  no  measurements  of  specimens  from  there.  Individuals 
from  Borneo  show  every  variation  between  the  two  extremes  of  colora- 
tion of  the  underparts  of  this  species.  Some  birds  are  paler  gray  on 
the  throat  and  breast  with  more  white  on  the  lower  abdomen.  Others 
are  darker  gray  with  less  white  on  the  abdomen.  Northern  Malay 
Peninsula  birds  seem  to  be  the  darkest,  while  Java  birds  are  the 
palest.  In  between  them  is  every  variation  of  what  is,  after  all,  a 
rather  slight  range  of  variation.  There  is  no  difference  in  size.  Under 
these  circumstances  it  seems  to  me  that  it  would  be  wiser  either  to 
recognize  several  microscopic  degrees  of  variation  and  name  all  the 
small  local  populations,  or  else,  and  this  I  think  preferable,  to  lump  all 
these  birds  as  longipennis.  This  would  leave  the  species  arranged  as 
follows : 

Hemiprocne  longipennis  coronata  (Tickell) 
India,  Ceylon,  Siam  to  Indo-China.    A  distinctive  rufous-throated 
race. 

Hemiprocne  longi pennis  longipennis  (Rafinesque) 
Peninsular  Siam  to  Sumatra,  Borneo,  Java,  and  Bali.    Dark  to 
paler  gray  below. 

Hemiprocne  longipennis  perlonga  (Richmond) 
West  Sumatra  islands.   A  large  race. 

Hemiprocne  longipennis  wallacii  (Gould) 
Celebes  to  the  Sula  islands.   More  bluish  on  the  upperparts. 


RIPLEY:   BIRD   FAUNA   OF   THE   WEST   SUMATRA    ISLANDS  355 

121.  Hemiprocne  comata  comata  (Temminck) 
Synonym:  Hemiprocne  comata  stresemanni  Neumann 

Nias,  Pini,  Tello,  Tana  Massa,  Tana  Bala,  Sibe'rut,  North  Pagi.  A 
series  from  these  islands  measures:  wing,  cf  126-130  (127.6),  9  123 
(worn)-134  (128.5).  I  fail  to  note  any  of  the  differences  mentioned 
by  Neumann. 

Family  TROGONIDAE,  Trogons 

Two  trogons  occur  on  the  west  Sumatra  islands.  One  from  Nias  is 
an  endemic  race  differing  slightly  from  its  nearest  relative  on  Sumatra 
and  the  Malay  Peninsula  in  color  and  a  tendency  to  have  a  larger  bill. 

122.  Harpactes  duvaucelii  (Temminck) 

The  only  record  for  this  species  is  Tana  Massa  (de  Schauensee  1940). 
A  male  and  a  female  taken  by  Kannegieter  measure:  wing,  c?  108, 
9  105. 

123.  Harpactes  oreskios  nias  de  Schauensee  and  Ripley 

Nias.  In  the  original  description  (1940,  p.  404.)  this  race  was  said 
to  differ  from  uniformis  by  lacking  the  orange  wash  on  the  chest  of  the 
male  and  having  narrower  white  bars  on  the  wing  coverts  and  second- 
aries. Three  males  and  a  female  from  Nias,  when  compared  with  birds 
from  Peninsular  Siam  and  the  Langkawi  islands,  do  not  show  these 
characters.  All  three  of  the  males  have  a  rich  orange  suffusion  indicat- 
ing that  the  type  may  not  be  fully  adult.  However,  these  Nias  birds 
do  have  somewhat  darker  more  brownish  crowns  than  the  birds  with 
which  they  have  been  compared.  Mr.  de  Schauensee  writes  me  (in 
litt.)  that  this  is  also  true  of  the  type.  As  well  as  this,  these  birds 
show  a  tendency  to  a  slightly  larger  bill.  The  culmen  of  the  four  Nias 
specimens  measures:  c?  17,  17.5,  18.5;  9  18.  A  series  of  uniformis 
measures:  c?  15,  15,  16,  16,  17;  9  14.5,  15,  16,  16,  17. 

This  trogon  is  found  in  heavy  secondary  jungle  at  low  heights  above 
the  ground,  often  in  substage  growth.  It  has  a  very  distinctive  bark- 
ing call. 

Family  ALCEDINIDAE,  Kingfishers 

The  west  Sumatra  islands  have  records  of  twelve  species  and  sub- 
species of  kingfishers.  Of  these  two  are  possibly  migrants  leaving  ten 
certain  resident  forms.  There  are  five  distinguishable  endemic  forms 
three  of  which  differ  primarily  by  size  and  two  of  which  differ  in  color 


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from  their  nearest  relatives.  Four  of  these  birds  are  most  closely  re- 
lated to  Sumatran  subspecies,  while  one,  Ceyx  erithacus  captus,  is 
almost  indistinguishable  from  C.  e.  motleyi  of  Borneo. 

Ramphalcyon  capensis 

This  is  an  interesting  and  widespread  species  ranging  from  Siam 
down  through  the  Greater  Sunda  islands  to  some  of  the  Lesser  Sundas. 
Oberholser  (Proc.  U.  S.  Nat.  Mus.,  35,  1909,  p.  657)  has  revised  the 
genus  and  ended  by  describing  several  races  of  capensis.  The  addi- 
tion of  further  specimens  to  the  National  Museum's  collection  has 
shown,  I  think,  that  this  action  was  unwise,  as  the  variation  in  plumage 
of  these  birds  is  considerable. 

At  least  in  the  Malay  Peninsula  and  island  part  of  its  range,  fresh 
plumage  in  this  species  is  assumed  in  July,  August,  or  early  September. 
At  this  time  the  color  of  the  pileum  may  approach  sepia  to  bistre. 
The  feathers  of  the  upper  back,  scapulars  and  wing  coverts  are  rather 
bright  greenish  blue  at  this  time.  After  eight  or  ten  months,  as  the 
feather  tips  become  frayed  and  the  color  of  the  pileum  bleached,  very 
noticeable  changes  are  brought  about  in  the  color  of  the  upper  parts. 
March  to  June  or  July  birds  have  the  pileum  colored  from  whitish- 
buff  or  pale  fawn  to  ochraceous-buff.  The  color  of  the  upper  back, 
scapulars  and  wing  coverts  changes  also,  this  time  not  due  to  bleach- 
ing, for  there  is  no  pigment  change  as  tests  under  sodium,  helium,  and 
ultra-violet  lights  have  shown.  Instead  the  frayed  and  sun-heated 
feather  seem  to  have  become  physically  changed,  contracting  the 
diffraction  grating  with  resulting  effects  similar  to  those  observed  in 
certain  beetles.  At  this  time  the  color  of  these  parts  varies  from  dull 
cerulean  to  indigo  to  purplish  blue. 

Birds  from  the  Malay  Peninsula  and  the  northern  part  of  Sumatra 
tend  to  be  somewhat  darker  buff  on  the  under  surface.  Perhaps  they 
should  be  kept  under  malaccensis.  Borneo,  Billiton,  and  Java  birds 
tending  to  be  rather  pale  beneath,  could,  it  seems  to  me,  quite  well 
be  lumped  together  as  capensis.  There  is  no  constant  difference  in 
size  in  a  series  of  measurements  of  more  than  thirty  specimens. 

From  these  birds  the  west  Sumatra  island  populations  differ  in 
being  somewhat  paler  and  more  washed  out  on  the  upper  surface  and 
by  a  tendency  to  larger  size.  The  blue  which  is  so  bright  in  capensis 
and  malaccensis  varies  in  these  specimens  from  dull  brownish  China- 
blue  in  a  September  female  from  Siberut,  to  pale  indigo  in  a  March 
male  from  Nias. 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  357 

124.  Ramphalcyon  capensis  simalurensis  (Richmond) 

Simalur.  The  type,  an  adult  male  collected  by  Abbott  November 
29,  measures:  wing  144.5,  tail  92.5,  bill  (from  naris)  07. 5.  Two  other 
males  and  three  females  measure:  wing,  cf  143.5  (molt),  143.5  (143.8); 
9  151,  155.5;  tail,  9  96,  99;  bill,  d1  65.5,  68.5;  9  70.5,  71.2. 

Stomach  contents  are  noted  as:  "small  fish,  prawns,  and  crabs." 
This  is  a  common  bird  of  the  mangroves  and  small  creeks  near  the  sea. 

Compared  to  malacccnsis,  this  series  is  equally  dark  buff  below,  but 
on  the  upper  surface  it  is  very  different,  lacking  the  bright  blue  or 
greenish  blue  of  the  upper  back,  scapulars,  and  wing  coverts. 

Compared  to  birds  from  the  other  west  Sumatra  islands,  these 
Simalur  birds  appear  uniformly  dark  buff  below  in  the  same  way  that 
malaccensis  differs  from  capensis.  The  upper  parts  also  appear  notice- 
ably duller  and  browner  than  the  majority  of  other  west  Sumatra 
island  birds,  but  this  condition  may  be  in  part  due  to  the  fact  that  all 
the  Simalur  birds  were  collected  between  October  23  and  November  30. 

125.  Ramphalcyon  capensis  sodalis  (Richmond) 

Synonym:  Ramphalcyon  capensis  nesoeca  Oberholser 
Synonym:  Ramphalcyon  capensis  isoptera  Oberholser 

Babi  (sight),  Tuangku,  Nias,  Pini,  Tello,  Siberut,Sipora, North  and 
South  Pagi.  The  type  of  sodalis,  an  adult  female  collected  January 
25,  measures:  wing  162,  tail  100.5,  bill  (from  naris)  73.5.  Another 
adult  female  from  Tuangku  measures:  wing  160,  tail  101.5,  bill  73.2. 
This  race  was  named  on  the  basis  of  size,  but  females  later  added  to 
the  Museum's  collection  from  other  islands  have  equalled  these 
measurements. 

The  type  of  nesoeca  from  Nias  is  an  adult  male  collected  March  15. 
It  measures :  wing  148.5,  tail  93,  bill  (from  naris)  68.5.  The  range  of 
this  race  was  given  (1909,  p.  674)  as  Nias  and  the  Batu  islands,  and 
the  characters  were:  "pileum  paler,  back,  wings,  and  tail  much  brighter 
and  more  bluish."  In  four  of  the  eight  specimens  this  is  true,  but  as 
these  are  March  birds  taken  at  a  time  when  the  pale  pileum  and 
brighter  blue  is  characteristic  of  the  state  of  wear  of  the  plumage,  I 
do  not  feel  50%  is  a  good  enough  average  difference  for  separation. 
However,  it  is  certainly  worth  noting  that  in  the  slightly  brighter 
color  of  the  upper  parts  these  four  Nias  specimens  indicate  a  closer 
affinity  with  Sumatra  than  is  the  case  on  the  other  islands. 

An  adult  male  taken  December  30  in  Sikakap  Strait  between  North 


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and  South  Pagi  is  the  type  of  isoptera.  It  measures:  wing  153.5,  tail 
94,  bill  (from  naris)  68.5.  This  race  was  described  (1909,  p.  671)  as 
being  larger  than  simalurensis  with  the  under  parts  lighter,  the  lower 
back  and  rump  more  greenish.  In  size  these  birds  agree  with  the  birds 
from  the  other  islands,  as  well  as  in  having  paler  underparts  than 
simalurensis.  I  can  find  no  constant  difference  in  the  greenish  char- 
acter of  the  rump. 

Measurements  of  a  series  of  sodalis  by  localities  follows : 


place 

wing 

tail 

bill  (from  naris) 

Tuangku  9 

160,  162 

100.5,  101.5 

73.2,  73.5 

Nias  c? 

146-152.5  (148.7) 

89-93  (91) 

68.2-72.5  (70.1) 

9 

163.5 

101 

68.5 

Batu  cf 

155 

89 

70 

O 

— 

92.5 

75 

Siberut  9 

156.5 

97.5 

71 

Sipora  cf 

150.5-158.5  (154.1) 

95.5-101  (98) 

66.5-70.5  (69) 

Pagi  c? 

150-153.5  (151.5) 

89-96.5  (93) 

63-70  (67.5) 

9 

153,  161 

91.75,  95 

68,  71.5 

126.  Alcedo  atthis  bengalensis  Gmelin 

Simalur,  Tuangku  (sight),  Nias,  Pini,  Siberut.  Two  pairs  from  Sim- 
alur  and  Nias  measure:  wing,  cf  70.5,  69.5;  9  70,  70.  These  birds 
were  taken  in  November,  February,  and  March. 

127.  Alcedo  meninting  meninting  Horsfield 
Synonym:  Alcedo  meninting  subviridis  Oberholser 
Synonym:  Alcedo  meninting  callima  Oberholser 
Synonym :  Alcedo  meninting  proxima  Richmond 

Simalur,  Tuangku,  Nias,  Pini,  Tana  Massa,  Tana  Bala,  Sipora, 
North  and  South  Pagi,  Enggano.  The  type  of  subviridis  is  a  male 
from  Nias  collected  March  23.  It  measures :  wing  62,  bill  (from  naris) 
34.  This  race  was  described  by  Oberholser  (1912)  as  smaller  than 
meninting  with  more  greenish  upper  parts.  Other  birds  from  Nias 
measure:  wing,  tf1  64.5,  9  63.5,  65.5;  bill,  cf  34,  9  31.5,  32.5.  Birds 
from  Sumatra  measure:  wing,  <?  61-65,  9  62-66.5;  bill,  c?  35-36.2, 
9  31.5-33.5.  Of  the  four  specimens  from  Nias,  only  one  is  especially 
greenish  on  the  back  and  rump. 

The  type  of  callima  is  a  male  collected  on  Tana  Bala  February  8, 
measuring:  wing  69,  bill  (from  naris)  36.2.  It  was  described  (1912)  as 
larger  than  meninting  with  upper  parts  slightly  more  greenish.  A  male 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA   ISLANDS  359 

from  Pini  and  a  female  from  Tana  Bala  measure :  wing,  d"  65,  9  68.5 ; 
bill,  cf  36.5,  9  34.5.  These  measurements  are  large,  but  Junge  (1936) 
gives  measurements  for  Simalur  birds:  wing,  d*  63,  66,  69;  9  65,  67, 
which  are  largely  overlapping.  Again  the  greenish  character  tends  to 
disappear  with  a  large  series. 

The  type  of  proxima  is  an  adult,  sex  undetermined,  from  North 
Pagi,  collected  January  4.  It  measures:  wing  69.5,  bill  37.5.  Other 
birds  from  North  and  South  Pagi  measure:  wing,  cf  67.5-69,  9  71.5; 
bill,  cf  35.5-37.5,  9  33.5.  This  race  was  described  as  being  very 
greenish  on  the  back  and  rump.  Two  of  the  specimens  are  particularly 
so — one  slightly,  and  the  fourth,  the  type,  can  be  matched  by  Sumatra 
birds.  I  think  this  character  is  entirely  too  variable  to  be  a  serviceable 
criterion  for  subspecies.  These  birds  are  also  large,  but  no  more  so 
than  Batu  or  Simalur  birds.  Recently  Junge  (1938)  has  recorded  a 
single  male  from  Enggano.   The  wing  measured  63. 

From  these  data,  I  would  be  inclined  to  think  that  probably  these 
kingfishers  at  some  time  or  other  reach  all  the  islands  of  the  group. 
Also  these  birds  show  a  tendency  to  larger  size,  particularly  on  Sim- 
alur, the  Batus,  and  the  Pagi  islands.  Finally  there  is  a  tendency  which 
I  do  not  consider  very  significant  so  far,  for  some  of  these  birds  to  have 
the  iridescent  back  and  rump  feathers  paler  and  more  greenish. 

128.  Ceyx  erithacus  captus  Ripley 

Nias.  This,  the  only  representative  of  the  species  from  the  west 
Sumatra  islands,  is  almost  identical  with  motleyi  of  Borneo.  From 
this  race  it  differs  by  a  longer  bill,  slightly  larger  size,  and  by  the  re- 
duction of  the  forehead  spot  which  may  be  lacking  in  some  specimens. 

A  not  uncommon  bird  in  thick  secondary  growth  or  substage  vege- 
tation usually  near  small  streams.  The  specimens  I  saw  were  all 
perched  about  six  feet  above  the  ground. 

129.  Ceyx  rufidorsus  jungei  Ripley 

Simalur,  Bangkaru  (sight),  Tana  Massa,  Tana  Bala.  These  birds 
differ  from  typical  ruflidorsus  by  larger  size.  A  series  measures :  wing, 
cf  62-64.5  (63.2),  9  62.5-63.5  (63);  tail,  <?  25-26  (25.5),  9  25.5-26.5 
(26);  bill,  cf  and  9  30.5-32  (31.3). 

130.  Ceyx  rufidorsus  rufidorsus  Strickland 

Siberut,  Sipora.  A  male  and  female  adult  and  an  immature  male, 
taken  October  28,  are  in  the  collection.   They  measure:  wing,  c?  61, 


360  bulletin:  museum  of  comparative  zoology 

9  62;  tail,  d*  25,  9  26;  bill  (from  naris),  tf  25,  9  26.  These  birds 
are  certainly  close  to  jungei  in  size,  but  Chasen  and  Kloss'  measure- 
ments (1926)  for  Siberut  and  Sipora  birds  are  so  small  that,  without 
seeing  the  rest  of  their  series,  I  would  rather  leave  them  for  the  time 
being  as  rufidorsus.  The  fact  that  one  of  the  specimens  in  the  National 
Museum  from  their  collection  is  a  subadult  male  inclines  me  to  be- 
lieve, however,  that  some  of  their  other  measurements  may  refer  to 
immature  specimens.  There  is  a  sight  record  for  Ceyx  from  the  Pagi 
islands,  which  may  refer  to  this  species. 

131.  Halcyon  coromanda  minor  (Temminck  and  Schlegel) 
Synonym:  Entomothera  coromanda  pagana  Oberholser 

Simalur,  Lasia,  Nias,  Tana  Massa,  Siberut,  North  Pagi.  The  type 
of  pagana,  an  adult  male  collected  January  4  on  North  Pagi  measures : 
wing  110,  tail  62,  bill  (from  naris)  44.5.  Three  females  from  Siberut 
and  North  Pagi  measure:  wing  107.5,  109.5,  110.5.  The  soft  parts  are 
listed  as:  "iris  dark,  sepia;  bill  and  feet  coral  red,  coral,  blood  red." 
These  birds  are  indistinguishable  from  minor  as  is  also  the  type  of 
neophora,  a  bird  from  Tapanuli  Bay,  northwest  Sumatra. 

132.  Halcyon  chloris  laubmanniana  Grote 

Synonym:  Halcyon  chloris  cyanescens  *  (Oberholser) 

Synonym:  Sauropatis  chloris  chloroptera  Oberholser 

Synonym:  Sauropatis  chloris  amphiryta  Oberholser 

Simalur,  Babi,  Nias,  Pini,  Lago,  Tello,  Tana  Massa,  Siberut.  Speci- 
mens from  this  area  measure  as  follows  (t  stands  for  type) : 


wing 

bill  (from  naris) 

Simalur  o" 

115,  116.5,  117.5  (t), 

117.5 

41,  43,  44.5  (t),  44.5  (2) 

9 

117 

46 

Nias  c? 

113.5  (t) 

42.5  (t) 

9 

114.5 

46 

O 

120 

43.5 

North  Pagi 

9 

111-116.5  (113) 

41-43.5  (41.8) 

The  type  of  cyanescens  from  east  Sumatra,  an  unsexed  bird  collected 
July  28,  measures:  wing  113,  bill  43.5.  Specimens  from  Sumatra, 
Banka,  and  Java  measure  up  to  115.5  (wing)  and  43.5  (bill).  The 
original  description  of  chloroptera  from  Simalur  (Proc.  U.  S.  Nat.  Mus., 
55,  1919,  p.  379)  compared  it  to  cyanescens,  but  said  the  birds  had 

1  Not  Halcyon  cyanescens  Cabanis  and  Reichenow,  1877. 


RIPLEY:    BIRD    FAUNA    OF   THE    WEST    SUMATRA    ISLANDS  361 

longer  wings  and  that  the  upperparts  were  more  greenish,  the  pileum 
was  darker,  the  blaek  nuchal  hand  narrower,  the  auriculars  less  green 
washed,  and  the  sides  and  flanks  noticeably  tinged  with  huff.  In 
series  these  color  characters  do  not  appear,  nor  are  the  size  measure- 
ments significant  when  the  amount  of  overlap  is  considered. 

In  the  original  description  of  amphiryta  from  Xias  (I.e.,  p.  382)  this 
bird  seems  to  stand  in  an  intermediate  position  between  chloroptera 
and  lavbmanniana.  It  is  smaller  than  the  former  and  larger  than  the 
latter.  In  color  it  is  brighter  than  the  former  and  duller  than  the 
latter.  This  is  a  form  of  microdissection  of  characters  which  escapes 
me.  The  plumage  changes  of  these  birds  are  now  well  enough  known 
to  make  comparisons  of  shades  of  greenish  or  bluish  color  untrust- 
worthy. Especially  is  this  difficult  when  a  series  of  birds,  for  example 
those  from  Simalur  (Oct.,  Nov.,  Dec.)  or  Nias  (Feb.,  March),  are 
taken  all  at  about  the  same  time.  Thus  they  may  present  a  condition 
.of  uniformity  of  plumage  which  is  not  really  valid  for  the  population 
as  a  whole. 

Undoubtedly  these  birds  will  eventually  be  shown  to  exist  on  all 
the  small  islands  and  reefs  of  this  group.  They  are  birds  of  the  man- 
groves and  beaches  and  usually  omnipresent  in  these  locations.  Junge 
(1936)  lists  nestlings  from  Simalur  collected  in  April  and  eggs  taken 
in  April  and  May.  Nesting  occurs  in  steep  banks  along  streams,  in 
hollow  trees,  and  even  in  termite  nests. 

133.  Halcyon  chloris  azela  (Oberholser) 

Enggano  and  Pulu  Dua.  This  race  resembles  birds  from  the  Malay 
Peninsula  and  outlying  islands,  the  Andamans,and  northeast  Sumatra, 
in  size,  but  on  the  average  tends  to  be  somewhat  darker  on  the  pileum 
and  more  dull  greenish  on  the  upper  back  and  scapulars.  In  view  of 
the  isolation  of  this  island  population,  it  seems  strange  that  these  birds 
are  so  slightly  distinct.  Actually  azela  appears  to  be  simply  a  small 
race  of  cyanescens,  like  that  race  somewhat  duller  and  darker  in  color 
than  the  small  Malay  Peninsula  birds. 

The  type  of  azela,  an  adult  male  collected  November  19,  measures: 
wing  103,  bill  (from  naris)  38.5.  A  series  measures :  wing,  c?  102  (worn) 
-104.5,  9  102-104.5;  bill,  c?  36-40.5,  9  38.5-41.5. 

The  races  arvistrongi,  humii,  and  davisoni  are  so  far  as  I  can  see, 
indistinguishable.  Sharpe  (Cat.  Birds  Brit.  Mus.,  17,  1892,  p.  277) 
describes  armstrongi  as  having  green  ear  coverts  and  the  black  band 
around  the  nape  obsolete.  This  description  does  not  seem  to  be  valid. 


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All  of  the  National  Museum's  series  from  Peninsular  Siam,  the  Malay 
Peninsula  and  islands,  the  Andamans,  and  Belawan  Deli,  Sumatra, 
have  at  least  partially  black  ear  coverts  and  a  well  indicated  nape 
band.  Robinson  (Birds  of  the  Malay  Penin.,  1,  1927,  p.  99)  felt  that 
the  kingfishers  from  Lower  Tenasserim  and  the  extreme  north  of  the 
Malay  Peninsula  were  more  dull  than  birds  from  elsewhere  and  so 
might  be  called  armstrongi.  The  National  Museum  has  both  dull  and 
bright  birds  from  all  localities,  which  inclines  me  to  believe  that  they 
should  all  be  listed  under  armstrongi,  the  name  with  page  priority. 

134.  Halcyon  pileata  (Boddaert) 

Simalur,  Babi  (sight),  Pini,  Siberut,  Sipora,  North  and  South  Pagi. 
This  species  has  not  been  found  breeding  south  of  Bangkok  and  may 
well  be  a  migrant.  Specimens  taken  in  November,  December,  and 
January  measure:  wing,  cf  130-131.5,  9  126.5-134.  Soft  parts  were 
recorded  as:  "iris  dark  brown,  bill  red  washed  with  brown,  feet  coral 
red."   The  stomach  of  one  bird  contained  a  lizard. 

135.  Halcyon  concreta  concreta  (Temminck) 

Siberut.  The  single  record  for  this  species  is  an  immature  male 
taken  by  Kloss.  The  wing  measurement  (1926)  is  recorded  as  112, 
and  the  soft  parts:  iris  dark,  bill  yellow,  culmen  black,  feet  greenish 
yellow. 

Family  MEROPIDAE,  Bee-eaters 

Two  species  have  been  recorded,  one  of  which  is  only  a  sight  record. 
One  species  which  is  a  migrant  has  been  collected  during  the  winter 
months. 

136.  Merops  leschenaulti  leschenaulti  Vieillot 

Dr.  Abbott  reported  seeing  this  species  on  Simalur  (Richmond, 
1903). 

138.  Merops  superciliosus  javanicus  Horsfield 

A  migrant  taken  on  Simalur,  Tuangku,  Nias,  Pini,  Tello,  Tana 
Massa,  Tana  Bala,  Siberut  and  South  Pagi.  The  occurence  in  these 
islands  is  from  September  19,  the  earliest  date,  to  March  16.  Some  of 
the  fall  birds  are  immature.  Their  plumage  is  worn,  the  cinnamon 
throat  patch  is  undeveloped,  and  the  upper  parts  are  sprinkled  with 
blue  tipped  feathers.    Stomach  contents,  winged  ants. 


RIPLEY:   BIRD    FAUNA    OF  THE   WEST    SUMATRA    ISLANDS  363 


Family  CORACIIDAE,  Rollers 

Two  forms,  both  migrants,  occur  among  the  islands  in  winter.  A 
third  resident  form  has  been  described  from  Simalur.  It  differs  from 
orientalis  of  the  Malay  Peninsula  and  Sumatra  primarily  in  color. 

139.  EURYSTOMUS  ORIENTALIS  ABUNDUS1  Ripley 

Simalur.  A  male  collected  by  Abbott  in  December  belongs  to  this 
race.  The  primaries  and  secondaries  are  washed  with  purplish  blue 
nearly  to  the  end  of  the  outer  webs.  The  specimen  measures:  wing 
194,  tail  99,  wing-tail  ratio  51,  wing  tip  index  36.  The  soft  parts  are 
recorded  as,  "iris  dark  brown,  feet  red". 

Records  from  Tello,  Sipora,  North  and  South  Pagi  may  refer  either 
to  this  or  the  following  race,  both  of  which  migrate  into  this  area. 

140.  EURYSTOMUS  ORIENTALIS  DEIGNANI  Ripley 

Nias.  A  male  collected  by  Abbott  in  March  presumably  belongs 
to  this  race,  although  in  worn  plumage  it  is  not  so  easy  to  distinguish 
the  color  characters  which  separate  it  from  abundus.  Certainly  there 
is  no  purplish-blue  wash  on  the  primaries  and  secondaries  and  the 
crown  is  rather  blackish.  Measurements:  wing  1ST,  tail  93  (worn), 
wing-tail  ratio  50,  wing  tip  index  40. 

141.   EURYSTOMUS  ORIENTALIS  OBERHOLSERI  Junge 

Simalur.  A  single  female  collected  in  October,  agrees  well  with 
Junge's  original  description.  It  measures,  wing  187.5,  tail  103,  wing- 
tail  ratio  54,  wing-tip  index  30.  Junge  (1936)  notes  that  females  were 
showing  nesting  activities  in  January  and  March.  Stomach  contents : 
large  hard  coleoptera. 


Family  BUCEROTIDAE,  Hornbills 

Two  species  occur  in  these  islands,  one  of  which  is  considered  to  be 
identical  with  Sumatra  birds  although  showing  an  incipient  tendency 
to  larger  size.  The  other  form  is  endemic,  differing  from  the  Sumatra 
and  Borneo  population  by  definitely  larger  size. 

1  replaces  Euryslomus  o.  catonyx  auctorum 


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142.  Anthracoceros  coronatus  convexus  (Temminck) 
Synonym:  Hydrocissa  convexa.  barussensis  Oberholser 

Nias,  Tello,  Tana  Massa,  Tana  Bala,  Siberut,  Sipora,  North  and 
South  Pagi.  The  type  of  barussensis  is  an  unsexed  bird,  obviously  a 
male  with  the  large  wing  measurement  of  326  and  a  tail  measuring  289. 
Birds  from  the  other  islands  measure  as  follows : 


wing 

tail 

Nias  d"  d1 

295-319.5  (306.7) 

269-294  (282.5) 

9 

275 

— 

Batu  Ids.  d1  d* 

314,  321 

285 

O  (=6) 

326 

289 

9  subad. 

279 

— 

Sipora  cT 

290 

— 

Pagi  Ids.  d*  & 

282-299  (289.6) 

273-282  (277.6) 

0(=    9) 

263 

246 

Birds  from  the  Batu  Islands  are  indeed  larger  than  average  but  I 
feel  that  there  is  too  much  overlap  to  make  this  race  recognizable. 
Measurements  of  over  300  for  the  wings  of  Sumatra  birds  are  not 
uncommon,  and  as  can  be  seen  from  the  above  measurements  there  is 
a  large  degree  of  overlap  between  Nias  and  Batu  birds.  However,these 
west  Sumatra  Island  birds  do  demonstrate  in  an  incipient  form  a 
speciation  response  by  increased  size  which  seems  to  be  so  character- 
istic of  these  island  bird  populations. 

Soft  parts;  iris  brown,  pale  vandyke  brown;  orbital  skin  bluish 
white;  bill  ivory,  patch  on  casque  and  base  of  bill  black;  feet  dull 
plumbeous. 

Weight:  &  2  lbs.,  9  1  lb.,  15  oz. 

This  is  a  rather  shy  bird  and  wherever  lumbering  or  gardening  tends 
to  reduce  the  original  forest,  they  become  much  reduced  in  numbers. 
In  the  south  part  of  Nias  where  there  is  little  original  forest  left,  they 
are  now  quite  scarce. 

It  is  perhaps  worth  noting  that  Hydrocissa  convexa  zamelaena  Ober- 
holser of  the  North  Natuna  Ids.  based  like  barussensis  on  larger  size 
appears  to  be  a  synonym  of  convexus.  The  type  measures:  wing  312; 
tail  308.  This  seems  to  be  another  case  of  a  small  island  population 
tending  towards  larger  size  than  a  mainland  or  large  island  population. 

143.  Crannorhinus  leucocephalus  megistus  Oberholser 

Tana  Massa,  Tana  Bala.  The  type,  an  adult  male  collected  February 
1 1 ,  on  Tana  Bala  measures :  wing  433,  tail  283.  Another  male  collected 


RIPLEY:    BIRD    FAUNA    OF  THE   WEST   SUMATRA    ISLANDS  3()5 

in  the  same  month  has  a  wing  of  408,  while  a  female  from  Tana  Massa 
measures;  wing  381,  tail  251.  I  have  compared  these  measurements 
with  ten  males  and  three  females  from  Pulu  Payong,  East  Sumatra, 
and  west  and  southwest  Borneo.  They  measure:  wing  cT  352-405 
(385.2),  9  347,  347,  354.  The  largest  Borneo  bird  measures  401.5. 
Comparing  these  measurements  by  use  of  the  formula  of  "t"  for  small 
samples  (Simpson  and  Roe,  Quantitative  Zoology,  1939,  p.  211)  I 
arrived  at  a  figure  of  3.07  for  the  measurements  of  the  males  and  8+ 
for  the  females.  Thus  the  probability  for  the  samples  being  from  the 
same  population  is  about  01.3%  showing  that  the  difference  is  signifi- 
cant. More  specimens  from  Sumatra  may  show  that  larger  birds  occur 
there,  but  for  the  present  I  feel  that  megistus  should  be  recognized. 

Soft  parts  are  recorded  as  follows:  cf  feet  blackish  green,  soles 
dirty  yellow;  cf  im.  iris  brownish  yellow,  feet  deep  dull  green,  bill  pale 
blood  red  at  base  shading  into  dull  greenish  yellow  at  tip;  9  bill  gen- 
erally yellow,  pale  toward  tip,  mottled  at  base  with  dull  brownish 
green.  Skin  of  throat  deep  dull  slaty  blue  with  a  few  pale  blotches, 
orbital  skin  paler  than  throat,  feet  grayish  olive. 

Weight,  c?  3.5  lbs. 

I  have  put  this  race  in  the  species  leucocephalus  as  I  consider  Ma- 
layan, Philippine,  and  Celebes  birds  conspecific.  Malayan  males  differ 
by  having  the  nape  black  instead  of  rufous  chestnut  but  in  two  birds 
of  the  National  Museum's  series  this  color  is  not  pure  black.  A  number 
of  small  brown  feathers  are  interspersed  along  the  margins  of  the  nape 
patch.  The  shape  of  the  bill  is  exactly  similar  although  the  color  is 
somewhat  different.  As  leucocephalus  is  the  oldest  name  I  should  list 
the  following  well  marked  races : 

Crannorhinus  leucocephalus  corrugatus  (Temminck) 
Malay  Peninsula,  Sumatra,  Borneo. 

C.  I.  megistus  Oberholser 
Batu  Ids. 

C.  I.  waldeni  Sharpe 
Guimaras,  Negros,  Panay. 

C.  I.  leucocephalus  (Vieillot) 
Camiguin  I.,  Mindanao. 

C.  I.  cassidix  (Temminck) 
Celebes. 


366  bulletin:  museum  of  comparative  zoology 


Family  CAPITONIDAE,  Barbets 

Two  endemic  forms  are  confined  to  the  west  Sumatra  islands. 
Both  differ  from  their  closest  relatives  on  Sumatra  primarily  in  having 
larger  bills  with  the  addition  of  certain  color  characters. 


144.  Chotorea  mystacophanes  ampala  Oberholser 

Tana  Bala,  Tana  Massa.  The  type,  an  adult  male  collected  February 
11,  1903  by  Dr.  Abbott  measures:  wing  104,  tail  59.5,  culmen  38.5. 
Two  males  and  two  females  measure:  wing  cf  99.5,  102.5,  9  102, 
105.5;  tail  &  52,  60,  9  55.5,  56.5;  culmen  &  37,  38.5,  9  37.5,  39.5. 

The  female  with  the  smaller  measurements,  collected  February  28, 
is  not  fully  adult  as  it  lacks  the  blue  tinted  throat  of  maturity.  Other- 
wise, except  for  a  slight  reduction  in  size  of  the  red  patch  on  the 
crown,  the  plumage  is  complete  and  must  be  presumed  to  be  that 
assumed  after  the  post ju venal  molt. 

Soft  parts :  feet  olive. 

This  race  is  chiefly  distinguished  from  typical  mystacophanes  by  the 
large  bill.  Both  males  and  females  also  seem  to  have  a  somewhat 
larger  red  crown  patch,  and  in  the  female  the  forehead  is  more  tinted 
with  blue  than  in  the  females  from  Sumatra  and  the  Malay  Peninsula. 
Birds  from  Peninsular  Siam  seem  to  show  a  slight  decrease  in  size 
indicating  that  there  is  a  continuous  cline  here  culminating  in  the  large 
Batu  Island  birds. 

In  spite  of  Chasen  and  Kloss'  remarks  (Treubia,  14,  1932,  p.  14.) 
I  cannot  see  the  validity  of  the  race  humei  from  Borneo 


145.  Cyanops  australis  gigantorhina  (Oberholser) 

Nias.  The  type,  an  adult  male  collected  March  26,  1903  measures: 
wing  74.5,  tail  38.5,  culmen  20.5.  A  series  of  four  males  and  three 
females  measure:  wing  c?  73.5-81  (76.5),  9  74.5,  75,  80.5;  tail  cf 
38.5-41  (40),  9  37,  38.5,  42.5;  culmen  &  19-20.5  (20.1),  9  20.5,  21, 
21.5.  Soft  parts  are  marked  as:  bill  black  with  gray  base,  black;  skin 
of  throat  dull  black;  feet  olive,  green. 

This  race  is  characterized  principally  by  the  size  of  the  bill  which  is 
noticeably  larger  than  in  Sumatra  birds,  four  males  and  three  females 
of  which  measure:  culmen  &  17-17.5  (17.2)  9  18-18.5  (18.3).  The 
color  of  the  underparts  is  slightly  variable  but  tends  in  Nias  birds  to 
be  a  purer  green  especially  in  the  males. 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  367 

Family  PICIDAE,  Woodpeckers 

Eleven  forms  occur  on  these  islands  of  which  nine  are  considered  to 
be  endemic.  All  nine  are  related  to  forms  found  on  Sumatra.  Of  them 
six  differ  primarily  by  color  characters  and  three  by  size.  Two  of  the 
size  races  are  larger  than  their  congeners,  one  is  smaller.  Of  the  two 
forms  considered  conspecific  with  their  Sumatran  relatives,  one, 
Dryobates  hardwickii  subsp.  shows  a  slight  inclination  towards  larger 
size  measurements. 

146.  Picus  puniceus  soligae  de  Schauensee  and  Ripley 

Xias.  This  race  differs  from  observandus  of  the  Malay  Peninsula, 
Sumatra,  and  Borneo  by  the  greatly  reduced  amount  of  yellow  on  the 
prolonged  occipital  crest  feathers.  In  the  type  and  two  specimens  of 
the  series  in  the  National  Museum,  yellow  is  quite  absent.  In  the  five 
other  specimens  in  the  National  Museum's  series,  yellow  is  present, 
but  it  is  overlaid  by  the  scarlet  vermilion  feathers  of  the  crown.  Three 
males  and  four  females  measure:  wing  d71  126,  126.5,  130,  9  126,  128, 
130.5,  132.5. 

Soft  parts :  iris  deep  crimson,  red  orbital  skin  pale  blue,  slaty  blue, 
leaden  blue;  bill  black  above,  ochraceous  beneath,  lower  ochre  yellow, 
yellow;  feet  dirty  yellow,  claws  horn  brown.   Weight  3.4  oz. 

As  Mayr  points  out  (Bull.  Raffles  Mus.  No.  14,  1938,  p.  30.)  birds 
from  Borneo  average  somewhat  smaller  than  Sumatra  or  Malay 
Peninsula  birds.  Two  females  and  an  immature  female  from  Banka 
are  larger  on  the  average  than  typical  observandus.  They  measure: 
wing  9  132.5,  136.5,  9  im.  138. 

This  is  a  bird  of  rather  thick  jungle,  often  found  in  dense  scrub 
growth  rather  close  to  the  ground. 

147.  Callolophus  miniaceus  niasense  (Buttikofer) 

Nias.  A  male,  three  females  and  one  unsexed  measure :  wing  o"  124, 
9  123,  123.5,  124.5,  o  125.5.  Birds  from  the  Malay  Peninsula  and 
Sumatra  measure:  wing  tf1  126,  127,  130,  9  126,  126,  129,  130. 

As  well  as  being  slightly  smaller,  this  race  seems  to  be  more  brightly 
red  on  the  crown  and  scapulars. 

148.  Dryobates  hardwickii  subspec. 

Nias.  A  single  male  collected  March  23,  measures:  wing  75,  tail 
33.5,  culmen  17.    The  latter  measurement  is  larger  than  a  series  of 


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birds  from  Java,  Sumatra  and  Borneo  which  measure:  culmen   cf 
12.5-16  (14.6)  9   13-16.5  (15.0). 

A  solitary  specimen  like  this  would  ordinarily  have  no  significance 
in  itself,  but  in  the  case  of  a  west  Sumatra  island  bird,  this  slightly 
larger  bill  does  point  I  think  to  a  potential  speciation  effect  in  this 
species.  Further  specimens  may  show  such  to  be  the  case.  Perhaps 
this  is  the  form  recorded  by  Nieuwenhuisen  and  Rosenberg  as  Picus 
percussus  (1868). 

149.  Meiglyptes  tristis  micropterus  Hesse 
Synonym:  Meiglyptes  grammithorax  microterus  Oberholser 

Nias.  A  series  of  five  males  and  one  female  from  Nias  measure: 
wing  90.5-94.5,  9  89.  Two  males  and  three  females  from  Sumatra 
measure:  wing  cf  93,  98,  9  91,  92.5,  95.  Oberholser 's  original  brief 
description  (1912,  p.  6)  of  microterus  was  "smaller".  Use  of  formulas 
(Simpson  and  Roe,  I.e.  1939)  shows  that  the  probability  of  these 
specimens  being  drawn  from  the  same  sample  is  greater  than  .1% 
Thus  these  small  differences  lack  any  significance.  In  color  and  mark- 
ing these  birds  are  identical. 

150.  Meiglyptes  tukki  calceuticus  Oberholser 

Tuangku.  This  race  was  described  by  Oberholser  (1912,  p.  6)  as, 
"like  Meiglyptes  tukki  tukki,  but  much  larger".  The  type  and  only 
specimen,  an  adult  female  collected  January  23,  1902,  measures:  wing 
112.2,  tail  70,  culmen  25.  Aside  from  size  there  are  no  other  significant 
differences. 

151.  Meiglyptes  tukki  infuscatus  Salvador! 
Synonym :  Meiglyptes  tukki  hylodromus  Oberholser 

Nias.  The  type  of  hylodromus  an  adult  male  taken  March  10,  from 
Nias  measures:  wing  97.5,  culmen  22.  It  was  described  without  ref- 
erence to  Salvadori's  race.  The  rest  of  the  series,  five  males  and  four 
females  measure:  wing  c?  95.5-103  (98.7),  9  91.5-100.5  (97.3), 
culmen  c?  22-24,  9  21-21.5.  These  measurements  agree  well  with 
typical  tukki. 

This  race  differs  from  tukki  of  Sumatra  by  having  less  well  defined 
pale  bars  on  the  back  and  scapulars  and  by  having  the  barring  on  the 
under  parts  more  reduced  on  the  average. 

A  male  of  tukki  from  Billiton  I.,  southeast  of  Sumatra  (U.S.N.M. 


RIPLEY:    BIRD   FAUNA    OF  THE    WEST    SUMATRA    ISLANDS  369 

180492)  seems  to  be  an  immature  bird.  Like  the  young  of  Dryobates 
the  feathers  of  the  crown  are  tipped  with  a  reddish  color,  in  this  case 
scarlet.  Another  mark  of  immaturity  may  be  the  pale  color  of  the 
abdomen  which  lacks  the  characteristic  dark  rusty  wash  of  the  other 
specimens. 

152.  Meiglyptes  tukki  batu  de  Schauensee  and  Ripley 

Pini,  Tana  Massa.  A  single  male  in  the  National  Museum's  col- 
lection has  a  wing  measurement  of  109.  It  differs  from  the  type  of 
calceuticus  by  having  the  barring  on  the  throat  more  broad  and  coarse, 
and  by  having  the  chest  more  blackish.  The  series  measured  by  de 
Schauensee  (1940)  seems  to  indicate  that  batu  is  intermediate  in  size 
between  tukki  and  calceuticus. 

153.  Micro,pternus  brachyurus  celaenephis  Oberholser 

Nias.  The  type  is  an  adult  female  taken  March  29,  1902  measuring: 
wing  109.5,  culmen  21.5.  Six  females  of  badius  from  Sumatra,  Banka, 
and  Billiton  measure:  wing  110-122  (113.9). 

As  may  be  seen  from  the  above  figures,  this  race  does  not  differ  from 
badius  by  being  "somewhat  larger",  (1912,  p.  6).  It  does  differ,  how- 
ever, by  being  "darker",  although  its  distinctive  characters  are  as 
follows :  crown  with  distinct  dark  centers  to  the  feathers,  upper  parts 
with  heavier  darker  bars;  under  parts,  breast  with  brownish  black 
subterminal  spots,  stomach,  flanks,  vent  and  under  tail  coverts 
heavily  barred  with  dark  brownish  black. 

154.  Dryocopus  javensis  parvus  (Richmond) 

Simalur.  The  type,  an  adult  male  collected  December  3,  1901  by 
Dr.  Abbott,  measures:  wing  185,  tail  124,  culmen  38.5.  A  series  of 
six  males  and  five  females  measure:  wing  d1  172-185  (177.7),  9  172- 
182  (177.2). 

Soft  parts;  iris  greenish  yellow,  straw  yellow;  bill  black;  feet  leaden. 

This  race  is  a  complete  miniature  of  typical  javensis. 

155.  Dryocopus  javensis  buttikoferi  (Richmond) 

Nias.  The  type,  an  adult  male  taken  March  18,  1903  measures: 
wing  232.5,  tail  167,  culmen  55.  An  adult  and  an  immature  female 
both  collected  March  25  measure  respectively;  wing  241,  215,  culmen 
53,  49.5. 


370       '      bulletin:  museum  of  comparative  zoology 

Soft  parts:  cf  iris  bright  yellow,  upper  mandible  black,  lower 
leaden,  feet  leaden;  im.  9  iris  straw  yellow,  feet  dusty  leaden. 

The  only  discernible  difference  between  this  race  and  javensis  is  the 
lack  of  the  blackish  bars  found  on  the  thighs  of  the  typical  race. 

I  doubt  if  these  birds  are  common  on  Nias  at  the  present  time.  The 
original  forest  on  the  island  is  not  plentiful  anywhere,  although  there 
is  more  of  it  at  the  north  end  of  the  island. 

156.  Sasia  abnormis  magnirostris  Hartert 

Nias.  Two  males  and  a  female  measure:  wing  cf  52.5,  53,  9  54; 
culmen  cf  14,  14.5,  9  15.  A  male  from  east  Sumatra  has  a  wing  of 
54  and  a  culmen  measuring  13.5,  and  eight  males  from  Borneo  and 
the  Malay  Peyinsula  measure:  culmen  12-13.5  (13.1).  Hartert  (1901) 
notes  that  his  birds  have  deeper  bills  at  the  base  but  I  cannot  find  this 
difference.  Computing  the  probability  of  these  culmen  measurements 
by  use  of  the  formula  of  "t"  (Simpson  and  Roe,  I.e.,  1939)  I  arrive  at 
a  figure  for  "t"  of  2.7  which  is  not  particularly  significant  being  more 
than  .02%.  However,  as  there  is  no  overlap,  I  believe  a  larger  series 
would  uphold  the  significance  of  these  measurements  especially  as  the 
measurements  of  four  females  from  Borneo  and  the  Malay  Peninsula 
are  13-13.5  (13.4),  giving  when  computed  by  the  "t"  formula,  a  prob- 
ability for  their  being  identical  of  only  a  very  little  more  than  .01%. 

Soft  parts:  iris  red;  ocular  area  purple,  lilac,  lilac-purple;  bill,  upper 
mandible  worn  brown,  lower  yellow;  feet  orange,  reddish  ochre. 

Weight:  cf  8.5,  11  grams,  9  11  gr. 

In  contrast  to  what  Robinson  says  (Birds  Malay  Penin.,  2,  1928, 
p.  115.)  I  found  this  bird  on  Nias,  not  in  deep  forest,  but  in  secondary 
scrub  near  cuttings  and  gardens.  In  June  a  male  with  enlarged  testes 
was  collected. 

Family  EURYLAIMIDAE,  Broadbills 

Four  forms  belonging  to  two  species  have  been  recorded  from  the 
west  Sumatra  islands.  Two  of  the  forms,  one  from  each  species,  are 
endemic,  differing  by  larger  size,  while  the  remaining  forms  are 
identical  with  Sumatra  birds. 

157.  Calyptomena  viridis  viridis  Raffles 

Nias,  Tana  Massa.  This  species  has  been  taken  by  Modigliani  and 
Kannegieter.   However,  it  has  not  been  met  with  since  their  time.   A 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  371 

single  male  from  Tana   Massa  recorded  by  de  Sehauensee  (1940) 
apparently  does  not  differ  from  the  typical  form. 

158.  Calyptomena  viridis  siberu  Chasen  and  Kloss 

Siberut,  North  and  South  Pagi.  A  series  measures  as  follows:  wing 
c?  107.5,  cT  im.  110,  111,  112  (2),  9  108.5,  112.5,  113.  Typical  viridis 
measures:  &  92.5-100.5,  9   98-106. 

In  size  this  race  agrees  with  continentis  from  Peninsular  Siam. 
However,  the  single  adult  male  is  darker  than  any  specimen  of  that 
form.  The  females  also  differ  from  females  of  continentis  by  being 
somewhat  darker  and  duller,  particularly  on  the  upper  parts. 

159.  Eurylaimus  ochromalus  mecistus  Oberholser 

Tuangku.  The  type,  an  adult  female  collected  January  29,  1902, 
measures:  wing  88,  culmen  19.  Two  other  females  measure:  wing 
87.5,  87.5,  culmen  18.5,  19.  These  birds  are  significantly  larger  than 
any  specimens  of  ochromalus  measured  by  me.  Apparently  they 
represent  an  isolated  population  on  Tuangku,  an  island  lying  not  more 
than  twenty-five  miles  off  the  west  Sumatra  coast. 

160.  Eurylaimus  ochromalus  ochromalus  Raffles 

Pini,  Tana  Massa.  Specimens  measured  by  de  Sehauensee  (1940) 
fall  within  the  range  of  ochromalus  rather  than  mecistus. 


wing 

culmen 

Batu  Ids.  (f 

79.5-83  (81.3) 

16.5-18. 

9 

79.5-84.5  (81.2) 

16-20. 

Malay  Penin.,  a1 

76-84.5  (80.7) 

16.5-20. 

Sumatra,  Borneo  9 

74.5-81.5  (78) 

16.5-18.5 

There  seems  to  be  no  difference  in  color  between  these  specimens. 

Family  PITTIDAE,  Pittas 

Two  pittas  occur  on  these  islands,  both  of  which  are  presumably 
migrants  rather  than  residents. 

161.  Pitta  brachyura  cyanoptera  Temminck 
Synonym:  Pitta  moluccensis  lepta  Oberholser 

Tuangku,  Nias,  Sipora.    The  type  of  lepta  is  a  male  adult  collected 
on  Nias,  March  15,  1903  .It  measures:  wing  121,  culmen  25.5.    A 


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male  and  female  from  Nias  and  Tuangku  taken  in  January  and  March 
measure:  wing  c?  121,  9  118,  culmen  cf  26.5,  9  25.  The  race  lepta 
was  described  as  being  smaller  than  moluccensis,  "especially  the  bill". 
A  series  of  males  from  Sumatra  measure:  wing  120.5-132.5,  culmen 
26.5-29. 

Aside  from  the  difference  in  size  being  insignificant,  Robinson's 
theory  ("Fasciculi  Malayensis,"  Zool.,  3,  1906,  p.  96.)  that  this  is 
a  migratory  form  rather  tends  to  mitigate  against  any  attempt  to 
delimit  isolated  populations. 

162.  Pitta  sordida  cucullata  Hartlaub 

Nias.  Another  migrant  which  has  been  recorded  only  by 
Buttikofer  (1896). 


Family  HIRUNDINIDAE,  Swallows 

Two  species  occur  on  the  west  Sumatra  islands.   One  is  a  migrant, 
the  other  conspecific  with  the  resident  form  of  the  greater  Sunda  area. 


163.    HlRUNDO  RUSTICA  GUTTURALIS  Scopoli 

Simalur,  Tuangku,  Nias,  Siberut.  A  migrant  recorded  from  the 
middle  of  September  to  the  beginning  of  May. 

164.  Hirundo  tahitica  javanica  Sparrman 
Synonym:  Hypurolepis  javanica  hypolampra  Oberholser 

Simalur,  Nias,  Pini,  Tello,  Enggano.  The  type  of  hypolampra  is  an 
adult  female  collected  on  Nias,  March  22,  1903.  It  measures:  wing 
106.  This  measurement  is  not  "larger"  than  typical  javanica  females 
which  range  from  103-108.  Nor  does  this  specimen  appear  strikingly 
pale  on  the  lower  parts. 

I  am  inclined  to  agree  with  Junge  (1936,  p.  46)  who  lumps  abbotti 
with  javanica,  as  there  seem  to  be  no  constant  size  or  color  differences 
between  the  populations  in  the  Greater  Sunda  area. 

This  is  a  common  bird  around  houses  and  villages.  It  often  rests 
under  the  eaves  of  European  houses.  Nesting  takes  place  from  early 
March  at  least  through  June.  This  species  will  probably  be  found  on 
all  the  islands  of  the  group. 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  373 

Family  CAMPEPHAGIDAE,  Cuckoo-shrikes 

Eleven  forms  have  been  recorded  from  these  islands.  Of  them, 
seven  are  endemic,  all  differing  from  the  Sumatra  forms  primarily  by 
size.  In  four  of  the  forms  there  are  minor  color  differences  which  set 
them  apart  one  from  the  other. 

165.  Coracina  striata  simalurensis  (Richmond) 

Simalur.  The  type,  an  adult  male  taken  November  19,  1901, 
measures:  wing  167.5,  culmen  27.5,  tail  120.5.  A  series  measures: 
wing  c?  168,  168.5,  169,  9  165,  165.5,  167.5,  9  im.  179;  culmen  d> 
29.5  (2),  30,  9  29,  29.5,  30,  9  im.  28.5. 

This  race  is  slightly  larger  than  sumatrensis.  The  males  are  slightly 
paler  gray  above  and  below.  The  females  are  paler  also  and  lack  the 
pronounced  barring  of  the  abdomen,  vent  and  under  tail  coverts  found 
in  sumatrensis.  Only  on  the  under  tail  coverts  are  the  bars  distinct. 
Elsewhere  they  are  virtually  obsolete.  The  immature  female  is  barred 
as  in  the  adults  of  sumatrensis,  but  the  bars  are  less  black  and  sharp  t 

166.  Coracina  striata  babiensis  (Richmond) 

Babi.  The  female  type  collected  January  13,  1902  measures:  wing 
172.5,  culmen  30.   Another  female  measures:  wing  172,  culmen  29. 

In  general  these  specimens  are  very  similar  to  simalurensis.  How- 
ever, the  barring  on  the  under  parts  is  even  more  reduced  than  in  the 
previous  race.  The  barring  on  the  under  tail  coverts  is  in  a  shade  of 
gray  on  white  rather  than  black  on  white.  In  color  this  race  approaches 
somewhat  the  tone  of  Coracina  papuensis,  a  mangrove  and  shoreline 
species  of  Australasia.  Presumably  the  habitat  of  babiensis  in  the  small 
low-lying  Banyak  Islands  is  rather  similar  to  that  of  the  widely  dis- 
tributed papuensis. 

167.  Coracina  striata  kannegieteri  (Buttikofer) 

Nias.  This  race  is  evidently  similar  to  sumatrensis  but  larger.  I 
have  seen  no  specimens. 

168.  Coracina  striata  sumatrensis  (S.  Miiller) 
Synonym:  Graucalus  crissalis  Salvadori 
Synonym:  Artamides  sumatrensis  halistephis  Oberholser 

Siberut,  Sipora,  North  and  South  Pagi.  I  cannot  see  any  difference 
between  these  birds  and  a  series  of  sumatrensis  from'Pulu  Tioman,  the 


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Rhio  Archipelago,  and  Great  Karimon  Island,  east  Sumatra.  The 
males  are  the  same  shade  of  dark  gray  on  the  upper  and  lower  parts. 
The  females  have  the  same  type  of  barring  on  the  underparts.  In  size 
there  is  too  much  overlap  to  provide  a  distinctive  character  for 
separation. 

Comparative  measurements  follow : 


wing 

culmen 

Siberut  d" 

161.5,  165.5 

26,  26.5 

9 

161 

26 

Sipora  c? 

169,  170.5,  172 

26,  27,  27 

9  (6) 

162-170  (166.4) 

26-28  (27.1) 

North  Pagi  d* 

— 

— 

9 

158 

28.5 

South  Pagi  &  (type 

!  of  halistephis) 

166 

25 

c? 

161 

26.5 

9 

162.5 

26.5 

Sumatra,  etc.  d* 

160-167.5  (164.1) 

27.5-29  (28.3) 

9 

158-168  (160.7) 

25.5-27.5  (26.6) 

169.  Coracina  striata  enganensis  (Salvadori) 

Enggano.  These  birds  are  very  close  to  simalurensis  both  in  size, 
which  varies  rather  widely,  and  in  color.  The  females,  however, 
differ  by  having  slightly  less  obsolete  barring  on  the  stomach  and 
vent.  Also  the  barring  on  the  under  tail  coverts  is  slightly  wider  and 
distinctly  more  blackish  than  in  simalurensis. 

Three  males,  four  females  and  an  immature  female  measure:  wing 
cf  166,  169,  173,  9  167,  167  (moult),  174.5,  179,  9  imm.  168;  culmen 
cf  28.5  (2),  30,  9  26,  27,  28,  9  im.  26.5. 

170.  Volvocivora  fimbriata  compta  (Richmond) 

Simalur.  Siberut  (?).  Two  specimens  were  collected  in  November 
and  December  by  Abbott.  The  type,  a  not  quite  adult  female,  measures : 
wing  103,  tail  83,  culmen  14.  Another  female  measures:  wing  100, 
tail  82.5,  culmen  14.5. 

Compared  to  females  of  culminata  this  race  is  larger.  Three  females 
from  Sumatra  and  Borneo  measure,  wing  92,  93,  93.5. 

A  male  from  Siberut  agrees  is  size  with  these  specimens  rather  than 
with  males  of  culminata  (wing  103  compared  to  92,93,  94.5,96  for  of  d* 
from  Sumatra  and  Borneo).  This  bird  does  not  seem  darker  than 
Sumatra  and  Borneo  birds,  but  lacking  males  of  compta  I  must  pro- 
visionally assign  it  to  this  race. 


RIPLEY:    BIRD   FAUNA    OF   THE   WEST   SUMATRA    ISLANDS  375 

171.  Lalage  nigra  nigra  (Forster) 
Synonym:  Lalage  nigra  empheris  Oberholser 

Nias.  The  type  of  empheris,  an  adult  male  collected  March  2, 
measures:  wing  88,  tail  6S,  culmen  15.  The  race  was  originally  based 
on  the  character  of  a  paler  rump,  but  I  cannot  find  that  the  type 
supports  this  contention. 

172.  Hemipus  hirundinaceus  (Temminck) 

Recorded  from  Simalur  by  Rosenberg  (1878)  and  from  Nias  by 
Xieuwenhuisen  and  Rosenberg  (1868). 

173.  Pericrocotus  flammeus  minythomelas  Oberholser 

Simalur.  The  type,  an  adult  male  collected  December  12,  1901 
measures:  wing  94,  tail  87,  culmen  14.5.  A  series  measures:  wing  d1 
91.5,  92,  93,  93.5,  94,  94,  9  89.5,  90,  91. 

This  seems  to  be  a  straight  size  race.  In  series  as  Junge  notes  (1938), 
the  females  only  show  color  differences,  not  the  males;  see  also  his 
1936  paper,  p.  53. 

174.  Pericrocotus  flammeus  modiglianii  Salvadori 

Enggano.  Fourteen  male  adults,  one  male  immature  and  four 
females  collected  on  Enggano  by  Abbott  agree  well  with  Junge's  dis- 
cussion (1938,  p.  350).  This  race  is  even  larger  than  minythomelas, 
wing  c?<?  93.5-97.5,  cT  im.  94.5,  9  9  92.5-94.5.  The  males  are  some- 
what more  yellowish  on  the  under  surface,  not  as  pure  vermilion  as  in 
xanthogaster.  The  color  of  the  underparts  of  the  females  is  inter- 
mediate between  xanthogaster  and  minythomelas,  while  the  upper  sur- 
face is  more  like  xanthogaster.  Except  for  the  fact  that  the  under  parts 
of  the  young  male  are  richer,  somewhat  more  orange-tinted,  particu- 
larly on  the  tail  feathers,  there  is  no  difference  between  this  plumage 
and  that  of  the  adult  female.  Specimens  collected  in  November  and 
December. 

176.  Pericrocotus  igneus  igneus  Blyth 

Nias.  A  single  record  in  Buttikofer  (1896)  is  the  only  one  for  this 
form  in  the  west  Sumatran  islands.  The  bird  was  a  female  taken  on 
north  Nias  in  the  fall  of  1895  and  is  listed  by  Junge  (I.e.  1936)  among 
his  material  of  the  nominate  race  which  he  compared  to  trophis. 


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Family  DICRURIDAE,  Drongos 

There  are  six  resident  drongos  on  these  islands.  Two  are  conspecific 
with  Sumatra  birds.  One  differs  by  size  from  a  Sumatra  race,  and  one 
differs  by  size  and  color  from  its  nearest  relative  on  Sumatra.  The  re- 
maining two  races  are  close  to  one  found  on  Borneo,  differing  from  it 
in  larger  bill  size  and  color. 


177.  Dicrurus  leucophaeus  leucophaeus  Vieillot 
Synonym:  Dicrurus  cineraceus  celaenus  Oberholser 

Simalur.  The  type  of  celaenus,  an  adult  male  collected  November 
27,  1901  measures:  wing  138.5,  tail  131.5,  culmen  22.5.  In  spite  of  the 
original  description  of  Oberholser  (1912,  p.  15),  "darker,  particularly 
on  lower  surface",  and  Junge's  supporting  view  (1936,  p.  62.),  I  cannot 
agree  that  the  Simalur  birds  differ  in  color  from  those  of  Java.  There 
is  no  distinct  difference  in  size  although  I  have  measured  no  Javan 
specimen  with  a  wing  as  long  as  138.  It  is  interesting  indeed  that  these 
birds  should  resemble  the  dark  Javan  race  rather  than  the  pale-lored 
Sumatran  forms  or  the  paler  white  eye-ringed  birds  from  the  islands 
farther  to  the  south. 

178.  Dicrurus  leucophaeus  siberu  Chasen  &  Kloss 

Siberut.  Four  males  and  one  female  of  this  race  measure:  wing  cf 
132,  135,  139,  141,  9  130.5;  culmen  c?  22-23.5,  9  23.5.  This  form 
has  somewhat  more  white  behind  the  eyes  and  is  a  trifle  paler  on  the 
back.  Otherwise,  except  for  its  larger  bill  it  is  very  close  to  stigmatops 
from  Borneo. 

The  appearance  of  this  white  eye-ringed  form  on  the  west  Sumatra 
islands  is  a  good  argument  for  including  the  former  species  lencogenis 
in  leucophaeus.  Presumably  this  color  pattern  is  carried  recessively 
in  the  uniform  gray  Sumatran  and  Javan  representatives. 

179.  Dicrurus  leucophaeus  periophthalmicus  (Salvadori) 
Synonym:  Dicrurus  leucogenis  diporus  Oberholser 

Sipora  and  the  Pagi  Islands.  In  the  original  description  of  diporus 
(1912,  p.  15)  Oberholser  made  no  mention  of  Salvadori's  race  from 
Sipora  with  which  Pagi  birds  are  identical.  The  type  of  diporus,  an 
adult  male  from  North  Pagi  collected  November  14,  1902,  measures: 


RIPLEY:    BIRD    FAUNA    OF   THE    WEST   SUMATRA    ISLANDS  377 

wing  142,  tail  127,  oilmen  24.  This  race  is  paler  than  siberu  with,  in 
most  specimens,  more  white  carried  back  from  the  eyes  and  down  onto 
the  sides  of  the  neck.  The  iris  is  marked  as,  "carmine",  "reddish". 
Moulting  birds  were  taken  in  September  and  October. 


ISO.    DlCRURUS  HOTTENTOTIUS  VIRIDINITENS  (Salvadori) 

Siberut,  Sipora,  and  the  Pagi  Islands.  A  series  of  specimens  have 
wing  measurements  as  follows:  cf  140  (worn)-152.5  (149.1)  9  139- 
148  (142.3).  The  iris  of  adult  birds  is  marked  as,  "deep  red"  or  "crim- 
son". An  immature  bird  taken  on  South  Pagi  in  September  has  the 
iris  marked  "brown".  Another  immature  was  taken  in  September  on 
Siberut.  See  Chasen  and  Kloss  (192G,  p.  293)  for  a  discussion  of  the 
differences  between  this  race  and  the  Sumatran  race. 


181.    DlSSEMURUS  PARADISEUS  PLATURUS  (Vieillot) 

Synonym:  Dissemurus  paradiseus  olizurus  Oberholser 
Synonym:  Dissemurus  paradiseus  pachistus  Oberholser 
Synonym:  Dissemurus  paradiseus  elassopterus  Oberholser 

Simalur,  Lasia  and  Babi.  The  type  of  olizurus  an  adult  male  from 
Simalur  taken  November  19,  1901,  measures:  wing  140,  tail  305, 
culmen  29.5.  The  type  of  pachistus  an  adult  male  from  Lasia  collected 
January  5,  1902,  measures:  wing  155.5,  tail  367,  culmen  30.  The  type 
of  elassopterus,  an  adult  male  collected  January  11,  1902  on  Babi, 
measures:  wing  154.5,  tail  imperfect,  culmen  31. 

Other  specimens  from  these  localities  have  the  following  measure- 
ments (excluding  the  tail  which  is  too  variable) : 


wing 

culmen 

Simalur  a" 

143 

29 

9 

134.5,  141,  141.5,  144.5 

26.5,  27,  27,  28 

Lasia  d1 

153.5  (2) 

27.5,  28 

Babi  d" 

147  (worn) 

30.5 

9 

144 

29.5 

I  agree  with  Junge  (1936,  p.  63.)  that  it  is  impossible  to  separate 
these  birds  from  the  Sumatran  platurus,  including  pachistus  which 
Junge  did  not  mention,  presumably  lacking  Lasia  specimens.  The 
worn  bird  was  collected  in  January. 


378  '  bulletin:  museum  of  comparative  zoology 

182.  Dissemurus  paradiseus  adelphus  Oberholser 

Nias.  The  type  of  this  race,  an  adult  male  collected  March  5,  1905, 
measures:  wing  157,  tail  347,  culmen  33.  A  small  series  measures: 
wing  <?  153.5,  157.5,  9  147.5,  148.5,  151;  culmen  d1  39.5,  31,  9  28, 
29.5. 

These  birds  are  larger  than  any  measured  by  Kloss  (Treubia,  13, 
1931,  p.  359.),  and  so  should  be  kept  as  a  separate  race.  Oberholser's 
other  characters  of  large  crest  feathers,  bristles,  racquet  feather,  etc., 
are,  I  feel,  not  particularly  diagnostic. 

Family  ORIOLIDAE,  Old  World  Orioles 

Four  subspecies  inhabit  the  west  Sumatra  islands,  three  of  which 
differ  from  the  Sumatra  form  by  larger  size  and  minor  color  characters. 
The  remaining  subspecies  is  based  on  color,  showing  close  Bornean 
affinities. 

183.  Oriolus  chinensis  mundus  Richmond 

Simalur.  The  type  is  an  adult  male  collected  November  19,  1901. 
It  measures:  wing  151,  tail  103.5,  culmen  35.  Four  other  males  meas- 
ure: wing  148-154  (150.7).  Besides  being  larger  than  maculatus  these 
birds  have  a  much  purer,  more  lemon,  yellow  colored  back. 

I  cannot  see  Chasen's  race  edgari  from  the  lower  Malay  Peninsula 
and  Sumatra.  A  female  from  north  west  Sumatra  and  another  from 
Banka  both  have  bill  measurements  (from  gape)  of  34,  thus  falling  well 
within  the  measurements  of  maculatus.  As  this  is  the  only  cited 
difference  (Treubia,  18,  Suppl.,  1941,  p.  118.),  I  fail  to  see  how  this 
race  can  be  maintained. 

There  is  a  record  for  this  species  from  Nias  in  Buttikofer  (1896). 

184.  Oriolus  chinensis  richmondi  Oberholser 
Synonym:  Oriolus  chinensis  siberu  Chasen  and  Kloss 

Siberut  and  the  Pagi  Islands.  The  type,  an  adult  male  collected  on 
North  Pagi  December  31,  1902,  measures:  wing  150,  tail  97.5,  culmen 
34.    Two  other  males  and  a  female  measure:  wing  c?  145,  152.5,   9 
148.  Birds  from  Siberut  measure:  wing  cf  149.5,  150.5,  152,  9  141.5, 
148.5. 

Chasen  and  Kloss  (1926,  p.  294)  described  siberu  as  being  larger 
and  greener  than  richmondi  without  citing  any  figures  to  support  their 


RIPLEY:    BIRD   FAUNA    OF  THE   WEST   SUMATRA   ISLANDS  379 

contention.  They  must  have  been  comparing  males  from  Siberut 
which  were  not  fully  adult.  A  single  male  in  the  U.  S.  National  Mu- 
seum's Siberut  series  is  fully  adult  and  is  indistinguishable  from 
richmondi.  It  is  quite  as  bright  orange  yellow  as  Pagi  birds  on  both 
upper  and  lower  surfaces.  Females  and  immature  birds  are  likewise 
inseparable. 

185.  Oriolus  chinensis  sipora  Chasen  and  Kloss 

Sipora.  Two  males  and  two  females  measure:  wing  c?  153  (2),  9 
148,  149.  These  birds  are  very  slightly  larger  than  richmondi  but  they 
differ  primarily  by  being  brighter  on  the  back  with  broader  yellow 
edgings  on  the  wings.  From  mundus  this  race  is  distinguished  by 
having  a  yellow  speculum  patch,  broader  yellow  margins  on  the 
secondaries,  and  by  being  somewhat  more  greenish  above.  It  is  curious 
that  this  race  should  occur  between  the  neighboring  islands  of  Siberut 
and  the  Pagi  group  where  richmondi  occurs.  The  inference  seems 
likely  that  Siberut  and  the  Pagi  Islands  are  inhabited  by  actual  geno- 
typically  separable  populations  which  have  the  misfortune  from  the 
taxonomic  point  of  view  of  being  phenotypically  indistinguishable. 

186.  Oriolus  xanthonotus  mentawi  Chasen  and  Kloss 

Siberut  and  Sipora.  Chasen  and  Kloss  neglected  to  compare  this 
form  in  their  original  description  (1926,  p.  295)  with  consobrinus  of 
Borneo.  The  single  adult  female  from  Siberut  in  the  National  Mu- 
seum's collection  is  inseparable  from  a  female  from  south-east  Borneo 
(U.S.N.M.  No.  181520.).  However,  the  two  adult  males  from  Siberut 
are  somewhat  more  boldly  streaked  on  the  breast  and  abdomen  than 
are  two  Borneo  birds. 

I  think  this  is  a  poor  race  which  might  be  shown  to  be  inseparable 
from  consobrinus  given  more  material.  One  of  the  females  of  the  latter 
form  from  the  Segah  river,  east  Borneo,  is  very  dark  gray  on  the 
throat  and  the  crown  has  broad  black  centers  to  the  feathers  making 
it  much  darker  than  any  other  specimen  examined. 

An  immature  male  was  collected  on  Sipora  in  October. 

Colors  of  adult:  "iris  crimson,  bill  pinkish  brown,  feet  grayish  black". 


Family  CORVIDAE,  CROWS 

Three  crows  occur  on  these  islands.    One  is  rare  throughout  the 
Greater  Sunda  area.    Of  the  other  two,  one  shows  a  slight  tendency 


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towards  a  larger  bill  compared  with  specimens  from  Sumatra  and 
Borneo,  while  the  other  form  is  identical  with  the  subspecies  found  on 
Java. 

187.  Corvus  enca  compilator  Richmond 

Simalur,  Nias.  Specimens  from  these  two  islands  have  somewhat 
larger  bills  than  Sumatran  and  Bornean  specimens,  but  I  have  not 
seen  a  large  enough  series  of  birds  from  the  latter  islands  to  determine 
whether  or  not  this  difference  is  constant.  Following  are  the  measure- 
ments of  seven  males  and  five  females  from  Simalur  and  Nias  as  con- 
trasted with  three  males  and  three  females  from  Sumatra  and  Borneo. 

wing  tail                      culmen 

Simalur,  Nias  d"          307-321.5(313.1)  146.5-170(157.5)  63-69.5(66.1) 

9     282  (\vorn)-310.5  (301.)  142.5-151  (146)  63-65.5  (63.8) 

Sumatra,  Borneo  <?             303-332  151-166  60.6, 62.5, 64 

9           303.5-318  148-169  59,  59.5,  63 

The  male  with  the  largest  bill  measurement  is  from  Simalur  and 
the  tip  of  the  bill  is  broken  as  is  that  of  another  male  from  Nias  with  a 
bill  measurement  of  66.5. 

In  color  these  birds  appear  to  be  slightly  more  blackish,  less  purplish 
on  the  under  surface  than  Borneo  and  Sumatra  birds.  The  iris  of  these 
birds  is  marked  as  dark  brown.   A  Nias  female  weighed  14.9  oz. 

This  bird  on  Nias  was  somewhat  shy  and  suspicious  of  humans.  I 
never  saw  it  out  on  the  reefs  or  beaches  where  orru  sometimes  goes. 

188.  Corvus  enca  enca  (Horsfield) 

Siberut,  Sipora.  A  small  series  from  these  islands  measures:  wing 
d1  276.5,  287.5;  9  273,  281,  293.5;  culmen  &  56,  59;  9  53,  54  (2). 
The  large-winged  female  is  unusual  in  this  respect  but  I  note  that 
Meinertzhagen  (Novit.  Zool.,  33,  1926,  p.  71.)  gives  wing  measure- 
ments for  this  form  up  to  299. 

A  female  collected  in  October  has  worn  primaries  and  newly  moulted 
rectrices.  A  name  on  one  of  the  labels  may  be  a  native  name :  "mengga". 

189.  Corvus  macrorhynchos  macrorhynchos  Wagler 

Recorded  from  Nias  by  Buttikofer  (1896,  p.  188).  This  species  is 
apparently  uncommon  in  the  area,  as  it  is  listed  by  Meinertzhagen 
(Novit.  Zool.,  33,  1926,  p.  85.)  as  "very  rare"  throughout  Sumatra. 


RIPLEY:   BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  381 


Family  SITTIDAE,  Nuthatches 

A  single  representative  of  this  family  has  been  recently  found  on 
Simalur.    It  does  not  appear  to  differ  from  the  Sumatran  subspecies. 

190.  Sitta  frontalis  saturatior  Hartert 

Simalur.  Junge  records  this  form  from  Simalur  (1936,  p.  60.)  where 
it  was  collected  for  the  first  time  by  Jacobson  and  van  Heurn.  He 
notes  that  it  was  locally  common,  sometimes  occurring  in  small  family 
flocks  in  high  open  forest  near  clearings. 


Family  TIMALIIDAE,  Babblers 

Eleven  babblers  occur  on  these  islands  of  which  five  belong  to 
endemic  races.  The  characters  which  separate  these  races  from  their 
nearest  relatives  on  Sumatra  are:  size,  two  forms;  color,  one  form; 
size  and  color,  two  forms. 

191.  Malacopteron  cinereum  niasense  (Riley) 

Nias.  The  type,  an  adult  male  taken  by  Abbott,  March  10,  1905, 
measures :  wing  80,  tail  64,  culmen  18.  Two  other  males,  a  female  and 
one  unsexed  measure:  wing  cf  80.5;  81.5,  9  73,  -  77.5;  culmen  d1 
16.5,  17,  o  17. 

This  is  a  distinctly  larger  race  than  that  from  the  Malay  Peninsula, 
Sumatra,  and  Borneo.  The  chestnut  forecrown  and  tail  are  also  some- 
what darker.  Soft  parts :  "iris  crimson,  reddish  brown ;  upper  mandible 
black,  lower  bluish  pink;  feet  bluish  pink,  pale  bluish  flesh".  Males 
with  enlarged  gonads  taken  in  June. 

As  pointed  out  by  de  Schauensee  and  myself  (1939,  p.  408.)  this  is 
the  species  listed  by  Salyadori  from  Nias,  not  magnum. 

192.  Malacopteron  affine  notatum  Richmond 

Bangkaru.  The  type,  an  adult  male  collected  January  17,  1902  by 
Abbott,  measures:  wing  80,  tail  68,  culmen  17.  Three  other  males  and 
a  female  measure:  wing  cf  75,  77,' 78,  9  73;  culmen  cf  15,  16,  17,  9 
15.5. 

Richmond's  original  description  (1902,  p.  190.)  mentions  that  this 
form  differs  from  affine  by  having  a  sooty  black  cap  instead  of  a  brown- 
ish one.   Actually  this  is  not  true.   Both  birds  have  a  sooty  black  cap 


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in  fresh  plumage,  but  notatum  is  somewhat  paler  on  the  upper  surface 
and  partially  lacks  the  dark  chest  band  characteristic  of  the  nominate 
race.  Also  it  is  larger  in  size  and  has  a  larger  bill.  Soft  parts:  "iris 
brown,  feet  leaden." 

193.  Malacopteron  albogulare  albogulare  (Blyth) 

Pini.  Two  specimens  listed  by  de  Schauensee  (1940,  p.  36.)  are  the 
only  records  for  the  west  Sumatra  islands.  His  remarks  indicate  that 
the  specimens  are  intermediate  between  albogulare  and  moultoni  but 
more  specimens  are  needed  in  order  to  determine  the  status  of  this 
population. 

194.  Anuropsis  malaccensis  malaccensis  (Hartlaub) 
Synonym:  Anuropsis  malaccensis  exsanguis  Oberholser 

Synonym:  Anuropsis  malaccensis  nesitis  Oberholser 

Tuangku,  Tana  Massa.  The  type  of  exsanguis,  an  adult  male  from 
Tuangku  collected  January  24,  1902  measures:  wing  72.5,  tail  35, 
culmen  17.5.  The  type  of  nesitis,  an  adult  male  from  Tana  Massa 
collected  February  20,  1903,  measures:  wing  77,  tail  36,  culmen  17.5. 
A  male,  a  female,  and  a  specimen  of  undetermined  sex  from  Tana 
Massa  measure:  wing  cf  78,  9  67.5,  o  69;  tail  c?  37;  culmen  d71  17, 
9  15,  o  15.5.  Sumatra  and  Malay  Peninsula  males  have  wings  meas- 
uring up  to  74,  and  tail  measurements  up  to  37.  The  culmen  measure- 
ments reach  17.  In  color  Tana  Massa  birds  are  very  slightly  richer 
more  rufescent  above  and  more  buffy  on  the  flanks  and  vent.  The 
Tuangku  specimen  is  exactly  intermediate  if  such  fine  distinctions  are 
possible.  I  concur  with  Riley  (unpublished  MSS.)  in  his  suppression 
of  exsanguis  but  I  should  like  to  carry  this  a  little  further  and  declare 
both  forms  synonyms  of  the  nominate  race.  I  feel  that  these  minor 
differences  in  size  and  color  are  too  overlapping  to  indicate  more  than 
a  trend  in  speciation  away  from  the  mainland  and  large  island  birds 
rather  than  a  positive  achievement. 

195.  Alcippe  brunneicauda,  brunneicauda  (Salvadori) 
Synonym:  Alcippe  cinerea  hypocneca  Oberholser 

Pini.  The  type  of  hypocneca  is  an  apparently  adult  bird  of  undeter- 
mined sex  collected  March  4,  1903.  It  measures:  wing  73,  tail  59, 
culmen  13.   A  male  also  collected  in  March  measures:  wing  70.5,  tail 


RIPLEY:    BIRD   FAUNA   OF  THE   WEST   SUMATRA    ISLANDS  •">*;! 

59.5,  culmen  13.5.  The  soft  parts  are  marked  as  iris  and  feet  grey,  bill 
brown.  In  measurements  a  series  of  Malay  Peninsula  and  Sumatra 
birds  completely  overlap  these  two  specimens,  so  that  they  can  hardly 
be  "smaller"  as  Oberholser  (1912,  p.  8)  diagnoses  this  race.  I  suggest 
further  that  the  fact  that  the  two  birds  are  in  rather  worn  plumage 
accounts  for  any  color  differences  that  may  be  present. 

196.  Stachyris  maculata  banjakensis  Richmond 

Tuangku.  The  type  is  an  adult  male  collected  January  24,  1902  by 
Dr.  Abbott.  It  measures:  wing  91,  tail  69,  culmen  21.  Another  male 
measures  wing  93,  tail  72,  culmen  20.  These  measurements  are  con- 
siderably larger  than  those  for  the  nominate  form  from  Sumatra:  wing 
cf  81,  84.5,  9  82;  tail  c?  67,  9  64;  culmen  c?  17.5,  20,  9  18.  Color 
differences  are  very  slight.  These  birds  are  somewhat  more  grayish  on 
the  upper  surface  but  two  specimens  is  not  a  large  enough  series  to 
establish  this  as  a  character. 


197.  Stachyris  maculata  hypopyrrha  Oberholser 

Pini,  Tana  Massa.  The  type  is  an  adult  male  collected  March  6, 
1903  on  Pini.  It  measures:  wing  85,  tail  65.5,  culmen  18.5.  This  small 
series  consisting  of  the  type,  a  female,  and  two  birds  of  undetermined 
sex,  agrees  well  with  Oberholser's  original  description  (1912,  p.  9)  in 
being  slightly  more  rusty  on  the  upper  surface  and  on  the  lower  flanks 
and  abdomen  than  a  series  of  three  birds  from  Sumatra.  There  is  no 
size  difference,  and  the  color  difference  is  so  slight  that  a  larger  series 
may  show  that  these  characters  are  not  constant. 

198.  Cyanoderma  erythroptera  fulviventris  Richmond 

Tuangku.  The  type,  an  adult  male  collected  February  1,  1902, 
measures:  wing  61,  tail  47,  culmen  17.  Another  male  and  two  females 
measure:  wing  cf  63,  9  59,  60;  culmen  cf  16.5,  9  15, 15.5.  Two  speci- 
mens from  Sumatra  measure;  wing  cf  (type  of  eripella)  60.5,  9  57; 
culmen  cf  14.5,  9  14.5. 

Besides  having  larger  bills,  these  birds  do  present  a  somewhat  more 
fulvous  appearance  on  the  upper  surface  and  on  the  flanks  and  lower 
abdomen.  Soft  parts  of  the  type  are:  iris  brownish  red,  throat  skin 
pale  blue.  The  feet  of  the  females  are  noted  as,  "pale  greenish  brown" 
and  "pale  brownish  fleshy". 


384  bulletin:  museum  of  comparative  zoology 

199.  Cyanoderma  erythroptera  erythroptera  (Blyth) 
Synonym:  Cyanoderma  erythroptera  pella  Oberholser 

Tana  Massa,  Tana  Bala.  The  type  of  pella,  an  adult  male  collected 
on  Tana  Massa  February  20,  1903,  measures:  wing  65,  tail  47,  culmen 
15.5.   Two  females  measure:  wing  58.5,  62;  culmen  14.5,  16. 

These  birds  are  fairly  large  but  the  measurements  indicate  a  good 
deal  of  variability  in  size. 

In  color  there  is  no  tenable  distinction  between  these  specimens  and 
erythroptera  with  which  they  were  not  compared  in  the  original  de- 
scription (1012,  p.  9).  The  soft  parts  of  one  of  the  females  are  given 
as  follows :  "orbital  skin  deep  cobalt,  throat  pale  whitish  blue,  feet  pale 
olive  yellow". 

200.  Mixornis  gularis  gularis  (Horsfield) 

Synonym:  Mixornis  gularis  zarhabdota  Oberholser 
Synonym:  Mixornis  gularis  zaptera  Oberholser 

Bangkaru,  Tana  Massa,  Tana  Bala.  The  type  of  zarhabdota,  an 
adult  male  from  Bangkaru  measures:  wing  61.5,  tail  52.5,  culmen  15. 
It  was  collected  by  Dr.  Abbott,  January  19,  1902.  The  type  of  zaptera, 
an  adult  male  collected  February  17,  1903  on  Tana  Massa  measures: 
wing  63,  tail  55,  culmen  15.5.  A  series  of  gularis  from  the  Malay 
Peninsula  and  Sumatra  measures:  cf  wing  58-64,  culmen  14-16. 

I  cannot  agree  with  de  Schauensee  (1940,  p.  37)  that  Batu  birds 
are  more  heavily  streaked  below  than  ordinary  gularis.  The  variation 
in  the  amount  and  type  of  streaking  is  considerable.  The  type  and 
unique  specimen  of  zarhabdota  is  rather  rufous  on  the  upper  surface 
but  this  condition  can  be  pretty  well  matched  by  Rhio  Island  birds 
and  must  be  considered  to  be  due  to  individual  variation.  Soft  parts 
are  indicated  as:  "skin  of  ocular  area  cobalt,  feet  yellow  olive". 

An  immature  female  was  taken  on  Tana  Massa  February  20,  1903. 
The  under  parts  are  grayish  buffy  except  for  the  throat  and  center 
of  the  breast  where  the  adult  yellow  feathers  with  central  shaft  streaks 
are  beginning  to  appear. 

201.  Macronus  ptilosus  ptilosus  Jardine  and  Selby 
Synonym:  Macronus  ptilosus  batuensis  Riley 

Tana  Bala.  The  type  of  batuensis  is  an  adult  male  taken  February 
13,  1903  by  Dr.  Abbott.  It  measures:  wing  73.5,  tail  65,  culmen  17. 
Another  specimen,  unsexed,  measures:  wing  71.5,  tail  59,  culmen  17.5. 


RIPLEY:   BIRD    FAUNA    OF  THE   WEST   SUMATRA   ISLANDS  385 

A  series  of  males  from  Sumatra  and  the  Malay  Peninsula  have  wing 
measurements  from  67-71.5  and  culmen  measurements  from  16-17.5 
so  that  I  do  not  feel  that  the  size  of  these  Batu  birds  is  particularly 
significant. 

Of  the  two  specimens  the  type  has  a  considerable  amount  of  gray 
on  the  abdomen  but  this  can  be  matched  by  individuals  in  the  series 
of  ptilosus.  I  am  inclined  to  think  that  the  amount  of  gray  in  this  area 
is  somewhat  due  to  the  makeup  of  the  skins. 

Soft  parts:  "skin  of  throat  deep  cobalt  verging  to  turquoise  on  neck 
and  eyelids". 

Family  PYCNONOTIDAE,  Bulbuls 

A  total  of  fifteen  members  of  this  family  have  been  collected  on  the 
west  Sumatra  islands.  Of  these  only  two  are  endemic,  although  five 
populations  which  are  considered  conspecific  with  those  of  Sumatra 
do  show  a  tendency  towards  larger  size.  Of  the  two  endemic  races 
both  show  slight  color  differences  compared  to  their  relatives  on 
Sumatra,  while  one  is  also  larger. 

202.  Aegithina  viridissima  viridissima  (Bonaparte) 
Synonym:  Aegithina  viridissima  nesiotica  Oberholser 

Tana  Massa,  Tana  Bala.  The  type  of  nesiotica  is  an  adult  male 
collected  on  Tana  Bala  February  5,  1903  by  Abbott.  It  measures: 
wing  62,  tail  42,  culmen  15.  I  agree  with  de  Schauensee  (1940,  p.  34.) 
that  this  is  not  a  tenable  race. 

203.  Aegithina  tiphia  horizoptera  Oberholser 

Nias.  The  type  is  an  adult  male  collected  on  Nias  March  3,  1905. 
It  measures :  wing  62.5,  tail  48.5,  culmen  16.5.  The  soft  parts  are  noted 
as:  "iris  whitish;  upper  mandible  black,  lower  blue  gray;  feet  blue 
gray."  As  was  pointed  out  by  de  Schauensee  and  myself  (Proc.  Acad. 
Nat.  Sci.  Phila.,  91,  1939,  p.  346.)  horizoptera  must  stand  as  the 
name  for  birds  from  Peninsular  Siam,  northern  Malay  Peninsula, 
Sumatra  and  Banka  as  well  as  Nias. 

204.  Chloropsis  sonnerati  parvirostris  Hartert 

Nias.  A  single  male  in  the  collection  of  the  Academy  of  Natural 
Sciences  measures:  wing  98,  tail  69,  culmen  22.5.  Its  weight  was  45 
grams  and  the  soft  parts  are  marked  as:  "iris  brown,  feet  gray". 


386  bulletin:  museum  of  comparative  zoology 

This  bird  was  in  breeding  condition  when  it  was  collected  (June 
10.).  It  was  perched  in  a  tall  tree  overlooking  a  cleared  garden  and 
was  singing  in  a  rich  burbling  manner  reminiscent  to  me,  of  the 
Baltimore  Oriole. 

205.  Chloropsis  sonnerati  zosterops  Vigors 

Pini,  Tana  Massa.  A  series  in  the  Academy  of  Natural  Sciences' 
collection  from  these  islands  measure:  wing  c?  101,  106,  106.5,  110, 
9  99.5,  100,  102.  This  is  larger  than  the  average  of  zosterops  from 
other  localities,  although  the  United  States  National  Museum  has 
specimens  from  Trang  reaching  104.  Certainly,  however,  the  Batu 
Island  birds  represent  a  tendency  towards  larger  size. 

206.  Irena  puella  criniger  Sharpe 
Synonym:  Glauconympha  cyanea  megacyanea  Oberholser 
Synonym:  Irena  puella  bondi  de  Schauensee 

Tuangku,  Nias,  Tana  Massa,  Tana  Bala,  Siberut,  Sipora,  South 
Pagi.  Oberholser's  type  of  megacyanea  is  an  adult  female  from 
Tuangku,  taken  January  23, 1902.  This  specimen  measures :  wing  121, 
tail  84,  culmen  25.  Soft  parts  are  noted  as  follows:  "iris  red,  bill  and 
feet  black." 

The  type  of  bondi  is  an  adult  male  (A.N.S.P.  no.  56496)  taken  on 
Tana  Massa,  which  measures:  wing  127,  tail  85,  culmen  25.  Fifteen 
males  from  these  islands  measure,  wing  121-134  (124.2)  while  eleven 
females  measure,  wing  115-128  (122.5).  Batu  birds  alone  measure: 
wing  c?  121-134  (124.1),  9  120-128  (123.9).  A  series  of  ten  males 
from  Borneo,  Banka  and  Sumatra  measure: